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VOLUME 69 NUMBER1 29 JUNE 2000 ;

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© The Natural History Museum, 2000

Entomology Series
ISSN 0968-0454 Vol. 69, No. 1, pp. 1-114

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London SW7 5BD Issued 29 June 2000

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Bull. nat. Hist. Mus. Lond. (Ent.) 69(1): 1—2 Issued 29 June 2000

EDITORIAL

This journal will soon celebrate its ‘golden jubilee’ (it first appeared as the Entomology Bulletin of the British
Museum (Natural History) in June 1950). Given the long history of both the Museum and taxonomy, it could as well
have started life in 1850 (and probably would have but for omnivores such as the Annals & Magazine of Natural
History). Looking at our Bulletin today, as I write within a few weeks of the millennium, I am forced to ask, what
sort of publication is this, and does it have a future? Is this aremnant of 20th century academic biology, or even 19th
century dilettantism, when colonels and men of the cloth were numerous amongst the greats of entomology? Is this
journal (and many others like it) simply struggling on, like one of those doomed lineages that just creeps past the
K-T boundary, only to disappear finally a few geological milliseconds later? Specifically, has the dawn of the web-
age now truly arrived, rendering print on white paper as outmoded for communicating science as tablets of stone
or scrolls of papyrus for chronicling human history? The answer is very assuredly no, but a qualified no. . .

Tasks of systematic entomology

Systematics, and systematic entomology in particular, is faced with a range of huge and seemingly unending tasks.
We still have the potential need to describe, at a very conservative estimate, three times the number of insect species
accounted for in the last 250 years. We need to deal with complex molecular, genetic, ontogenetic, polymorphic,
phenetic, seasonal and geographical variation, and all that this means for problems in synonymy. We must strive
to provide far better means of identification: even for those insects that have a more or less reliable name and
diagnosis, most can still only be named by experts working with large collections. We must strive to provide a
robust predictive classification: increasingly this means team-work to create enormous morphological and
molecular data sets. While progress with empirical cladistics is now impressive, the vast majority of insect species
(and even genera) are still either wholly unknown, or poorly sampled and little studied. Without sound predictive
classifications, our attempts to understand, conserve and make best use of the vast diversity of insects will remain
largely ineffectual.

The challenge that information technology brings

With the emergence of computer technology, including the ability to store, manipulate and transmit vast amounts of
data, as well as excellent digital images, all by internet, anew challenge has emerged for systematic entomologists.
In addition to all the basic functions of research (and the necessary curation of huge museum collections that
underpin our work), systematic entomologists are now also expected to act as information scientists and providers.
The world is no longer content with the idea that, somewhere in Washington, Canberra, Paris, Berlin, London,
Nairobi, Bangkok or Quito, there are obscure but incomplete lists and descriptions of various species and genera,
barely accessible even to those museum workers traditionally expected to create and care for them.

Instead, we are in the age of biological informatics. Such data should be ‘on the net’, there for all to have access,
to see what it says on the labels of all the Noctua pronuba, Schistocerca gregaria, Carabus violaceus, Aedes aegypti
oramillion other species in ten thousand museums, all available at the flick of a switch, the touch of an icon. Together
with on-line images of the specimens, fully synonymic lists of the groups to which they belong, interactive keys for
identification, and automated mapping programmes. Even better of course, rather than do any of this work yourself,
the relevant specialists are only an e-mail away, to receive new data, verify it, and feed it back into the maw of the
gargantuan, all-consuming internet machine that will link old museum samples, new field samples and users all
together, thus to receive the wisdom and blessing of the experts. Stirring stuff, politically correct too — but consider
those experts for a moment. What chance for reflection will they have, what chance to absorb impressions, form
ideas, create new information, create new science? Precious little, if we get the balance wrong. Or, if we get it right,
a wonderful broadening of their own horizons and a re-affirmation of their central role in biology.

If the relatively few insect taxonomists that exist are really expected to make their knowledge and expertise
available to all, then unless they receive very significant extra support, they will be overwhelmed. Equally, if the
professional community of systematists fails to respond to the challenge of the web, and respond well, then they
will be marginalised. What government would continue to support taxonomists incapable of making their work
available by the greatest new aid to communication in 500 years, a veritable technological revolution so favourable
and helpful to the basic tasks of taxonomy and systematics, if only it can be tamed and used wisely?

© The Natural History Museum, 2000

2 EDITORIAL
Pursuing and delivering basic research in taxonomy

The great threat that is implicit in the informatics imperative is that taxonomists will be pulled away from their
fundamental tasks of original research (creating new knowledge) and documentation (developing and caring for
the collections that record their work and provide the basis for future research), to focus on re-packaging and
dissemination of what is already known. Basic research in taxonomic entomology involves a massive and
unfinished task of discovery: the discovery of taxa (with a majority of species still to be found and recognised), and
the discovery of taxonomic characters (fundamental for the reconstruction of phylogenetic relationships as the
basis of predictive classifications). Related tasks include establishing the distribution and basic ecology of species,
application of formal names (zoological nomenclature), and understanding the significance and origins of diversity
(comparative and evolutionary biology).

While accepting that original research and the curation of collections are the sine qua non of systematic
entomology, we must also acknowledge that making discoveries accessible to humanity as a whole is also
fundamental to science. As discussed, the emergence of the internet can now be seen as a key means of delivering
our science to the widest possible audience. However, the internet is, as yet, still essentially ephemeral. Its very
existence is now dependent on events completely beyond the control of scientists. Moreover, the traditional
printed word on paper still has enormous appeal and practicality.

There is an emerging view, to which I subscribe, that the internet and the printed word have different strengths
and weaknesses, and that we should seek to exploit the strengths of both to make our work as accessible and useful
as possible. Different roles will develop for ‘hard copy’ and ‘electronic’. With serious new investment by The
Natural History Museum in electronic communication now underway, and a emerging commitment to make sure
that we play our full and proper role in this access revolution, I also look to this good but still relatively obscure
journal to take on a new lease of life, to take on a new role as we strive to find the way to make best use of these
wonderful media, literally in our hands and at our fingertips.

Rather than just fade away like an old dinosaur, I therefore expect this Bulletin (and other museum publications
like it) to emerge from the shadows, rather like those meek little mammals that supposedly lurked while
Tyrannosaurus roamed. However, I was reminded by a colleague earlier today that the weight of termites per
square metre of soil in a Cameroon rainforest is equivalent to the density of people on the streets of Tokyo or
London. Time to get back to the microscope I think, for fear that we will have nothing new to say in this brave multi-
media world. But the opportunity to communicate about our science and about our collections on a grand and more
effective scale than ever before is surely here, if we can but grasp it. I look forward to seeing how we and our
entomological colleagues around the world rise to these exciting challenges, confident that this Bulletin, like those
once obscure mammals, will evolve rapidly to take full advantage of the new opportunities, and play a key role in
the process. Long live the internet — and long live our vital ink on paper.

R. I. Vane-Wright

Keeper of Entomology

The Natural History Museum
London, Ist December 1999

Bull. nat. Hist. Mus. Lond. (Ent.) 69(1): 3-114 Issued 29 June 2000

A review of the Central and South |

American Nepticulidae | GHE NATURAL
(Lepidoptera) with special i fo oe ,
reference to Belize ;

PRESENTED

“sy

RIMANTAS PUPLESIS

yy Department of Zoology, Vilnius Pedagogical University, Studentu str., Vilnius 2034
"LT, Lithuania

GADEN S. ROBINSON
Department of Entomology, The Natural History Museum, Cromwell Road, London
SW7 SBD, UK
CONTENTS
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SyNopsIS. Fifty-eight species of Nepticulidae are described or redescribed from Central and
South America; 16 Neotropical species are redescribed, 33 are described as new and 4 are
described but not named pending the availability of additional material; five species from Florida,
considered to be of Neotropical affinity, are listed together with citations of descriptions and
illustrations. Four new combinations are established; one new species-group is defined in
Stigmella. The generic status of Manoneura Davis is restored. Almost all primary types have been
examined and illustrated for the first time except for some predominantly non-tropical species. The
species revised fall into seven genera: Enteucha (6 species), Manoneura (2 species), Stigmella (23
species), Ectoedemia (5 species), Fomoria (4 species), Acalyptris (16 species) and Glaucolepis (2
species). Adults of all species treated (except for a few from Florida) are illustrated, with line
drawings of genitalia and distribution maps.

© The Natural History Museum, 2000

4

INTRODUCTION

Committed effort to inventory the world’s biota has
never been more necessary than now. Growing inter-
national concern over the biodiversity crisis together
with the provisions of the 1992 Rio Convention on
Biological Diversity have imparted a new urgency to
the need for taxonomic revisions of diverse groups and
the provision of identification manuals, particularly
for tropical areas. Hence this publication.
Nepticulidae are a very specialized, isolated family
of primitive monotrysian Microlepidoptera with a
worldwide distribution. The minute size of the adults,
the concealed mining life-style of the larvae (predomi-
nantly in leaves), and the difficulty of rearing imagines
goes some way towards explaining why these moths
are still poorly studied in many regions. Only the
northern European nepticulid fauna can be considered
to have been exhaustively studied (Johansson et al.,
1990). Studies in other regions of the world fall some
way short of reflecting the actual diversity of the group
although coverage compares well with that of other
groups of Microlepidoptera. The most exhaustive revi-
sions or detailed taxonomic papers have been published
for the western Palaearctic (Johansson, 1971;
Nieukerken, 1983, 1985a, 1985b, 1986a; Johansson et
al., 1990; Nieukerken & Puplesis, 1991 etc.), Central
and Eastern Palaearctic, including Far-eastern Russia
and Japan (Kemperman & Wilkinson, 1985; Puplesis,
1994; Puplesis & Diskus, 1995, 1996a, 1996b, 1996c;
Puplesis et al, 1996, 1997), South Africa (Scoble,
1978a, 1978b, 1980a, 1980b, and, outstandingly sig-
nificant, 1983), Nearctic (Davis, 1978; Wilkinson,
1979, 1981; Wilkinson & Scoble, 1979; Wilkinson &
Newton, 1981; Newton & Wilkinson, 1982), New
Zealand (Donner & Wilkinson, 1989), and recently for
Australia (Hoare et al., 1997; Hoare, 2000). Long
neglected, the diverse Australian fauna is being further
studied by Robert Hoare; the Chinese and Japanese
fauna is being revised by Erik van Nieukerken (NNM);
arevision of the Central Asiatic nepticulids is currently
in preparation by R. Puplesis and A. Diskus (VPU).
The Neotropical Region, while vast and with a
hugely diverse biota, is comparatively unexplored with
respect to collection and study of Nepticulidae. The
first two species recorded from the region, Stigmella
Johannis and Fomoria molybditis, were described by
Zeller (1877) (as Nepticula) from Colombian material
collected by Johann and Nolcken in 1871. Much later,
five further species were reported from Peru and two
from Guyana by Meyrick (1915); however, one of the
Guyanese species (Enteucha cyanochlora) was placed
in the Lyonetiidae and only recognized as nepticulid
some 70 years later (Davis, 1984; 1985). During the
last seven to eight decades there have been practically
no investigations of Neotropical Nepticulidae, except

R. PUPLESIS AND G.S. ROBINSON

for isolated descriptions of a few new species from
Puerto Rico (Forbes & Leonard, 1930) and Argentina
(Meyrick, 1931; Bourquin, 1962) and a few relatively
recent discoveries in Florida (Davis, 1978; Wilkinson,
1981) of species with Neotropical affinities (Fig. 1).
Stigmella plumosetaeella was described by Newton &
Wilkinson (1982) from Arizona, USA, and here it is
additionally recorded from south-western Mexico.
However, until now only 21 species have been recorded
from Central and South America and the tropical states
of the USA. Thirteen of these have been listed by Davis
(1984) in the Neotropical checklist, but most have
never been illustrated.

In total fifty-eight species of Nepticulidae from
Central and South America are treated in the present
revision. We also include five species currently known
from southern Florida as potential members of the
Neotropical fauna; the practicality of this is confirmed
by recent discovery of ‘Florida species’ in Belize and
Dominica (e.g., Manoneura basidactyla). Another eight
species with a distribution range from northern Florida
into the northern states of the USA have not been
incorporated into our revision as their hostplant data
and distribution suggests they are typical members of
a temperate fauna; they are listed at the end of the
checklist below. The species revised here fall into
seven genera: Enteucha Meyrick (6 species),
Manoneura Davis (2 species), Stigmella Schrank (23
species), Ectoedemia Busck (5 species), Fomoria
Beirne (4 species), Acalyptris Meyrick (16 species)
and Glaucolepis Braun (2 species). Only Manoneura
is known exclusively from the Neotropics.

This revision is the first attempt to demonstrate to
the scientific community an example of the diversity of
Neotropical nepticulids and to prepare the ground for
future investigations of this fauna. Substantial uniden-
tified material was available to us, collected during the
last thirty years and deposited mainly in the University
of Copenhagen and in the United States National Mu-
seum of Natural History (Smithsonian Institution),
together with material that we have collected recently
in Belize.

The Belize collecting trip, undertaken in April 1998
by the first author with the assistance of Simon Hill
(University of Westminster, UK), has resulted in the
description of a very interesting and rich rainforest
fauna of nepticulids and other primitive
Microlepidoptera. The nepticulid species found in
Belize comprise exactly half of all species treated in
the present revision. Belize has given us a baseline
from which to estimate the possible extent of
Neotropical nepticuloid diversity, allowed us to com-
pare the overlap of the Belize collection with previous
samples, and to supplement the existing BMNH hold-
ings (otherwise almost entirely aged type material).
Nearly one half of the country visited lies in dense
protected forests, some still unexplored. Except for a

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

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Figs 2,3 Sites in Belize where fieldwork was conducted in April 1998. 2, site at Pook’s Hill Nature Reserve; 3, habitat
near Las Cuevas Research Station, Chiquibul Forest.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

few local collecting trips, two main sites were used:
the Las Cuevas Research Station and Pook’s Hill
Nature Reserve (Figs 2, 3). The Las Cuevas Station
was built recently and is operated jointly by the Gov-
ernment of Belize and The Natural History Museum,
London. It is situated deep in the hilly Maya Moun-
tains in the Chiquibul Forest, Cayo District. The
majority of the vegetation is intermediate-age (ca. 30
yrs) secondary forest, with the exception of a few
pockets of relatively undisturbed or primary forest.
The Chiquibul receives between 1500 and 2000 mm
of rain per annum. Rainfall is not distributed regularly
throughout the year for there are two dry seasons
(from February to June and again for two to three
weeks in August). Hilltops, hillsides, and valley bot-
toms have markedly different forest both in terms of
structure and species composition. At our main col-
lecting sites (on hillsides and valley bottoms) the
forest has a greater proportion of evergreen species,
e.g., Manilkara zapota, Calophyllum brasilense and
Aspidosperma megalocarpon, although deciduous
species such as Swietenia macrophylla and Terminalia
amazonia are still an important component. Pook’s
Hill is a 300-acre private Nature Reserve of pristine
jungle set in the foothills of the Maya Mountains,
surrounded with forest and bordered by one of Be-
lize’s largest and best-known nature reserves, the Tapir
Mountain Nature Reserve (see Puplesis & Hill, 1998).

Most of the material collected by Puplesis and Hill
was obtained by light trapping, with only a single
species reared from the larval stage. According to our
field observations and those of Dr Owen Lewis
(BMNH), who has spent a continuous year studying
leaf-miners at the Las Cuevas Station, mining larvae
are very sparse in April but Owen (pers. comm.) col-
lected numerous fresh mines (although not those of
Nepticulidae) in November—January.

The material from Belize has yielded twenty four
new species; another four species also appear to be
new but they are left unnamed mainly because of the
poor condition of the specimens available. Only one
species collected in Belize (Manoneura basidactyla
Davis) was previously known, from southern Florida
(and it is recorded additionally here from Dominica).
The taxonomic composition of the Belize nepticulid
fauna has a few very interesting features. It includes a
new species in the previously monobasic genus
Manoneura, considered by Nieukerken (1986b) to be a
synonym of Enteucha, but here restored to full generic
status. There are three morphologically very distinct
new species in Enteucha, a tiny genus comprising just
seven species, six from the Neotropics, four of which
are described in the present revision. One of the most
unexpected results of the Belize expedition was the
discovery of a diverse fauna of Acalyptris, a total of 14
species representing 48% of Nepticulidae recorded
from the area or about 19% of the known species of the

7

genus. The Belize Acalyptris exhibit a remarkable
range of morphological structure. The dominance of
Acalyptris might be a seasonal phenomenon, for the
Belize collection was made during a short period at the
height of the dry season when Microlepidoptera were
otherwise infrequent at light. The sparse sampling of
nepticulids in most of the Neotropical Region, despite
the vast potential of their probable habitat, suggests
that what we are seeing in Belize is just the tip of the
iceberg.

The pattern of diversity of the Belizean fauna may
be seen in the annotated checklist, below. The com-
paratively small number of species of Ectoedemia
(s. str.) is surprising but may just be the result of under-
sampling. However, only three species are known from
southern Africa (Scoble, 1978a). We have not recorded
representatives of Simplimorpha, Bohemannia,
Parafomoria, Trifurcula (s. str.) and Etainia from the
Neotropical region. These genera have mostly restricted
distribution ranges in the eastern hemisphere or Eura-
sia. The Australian endemic Pectinivalva and recently
described Roscidotoga have not been found in our
samples either.

We have noticed unusually strong and independent
reduction of wing venation among the species investi-
gated, and this limits the use of venational characters
for generic recognition in the American tropics. The
genital morphology of the newly discovered or newly
dissected Neotropical nepticulids includes a few inter-
esting features not known in other faunas (see, e.g.,
Stigmella ovata, Ectoedemia fuscivittata).

Species from the lowland tropical forest of Belize
seem morphologically more distinct and apparently
more isolated both from each other (and from boreal
species of the same genus) than are species from the
Andes and Patagonia. Species differences in the An-
dean fauna are generally slight, and problems of
identification of similar species are compounded by
intraspecific variation. In Stigmella epicosma, for
example, both forewing pattern and genital morphol-
ogy vary with the altitude of the habitat. The
preliminary impression of the neotropical nepticulid
fauna is one of two distinct faunal elements. The first
is a relatively young element of species with distinct
boreal affinities and showing some evidence of recent,
rapid speciation and localized geographical variation,
occurring in the Andes and the southern temperate
zone. The second element comprises more clearly
differentiated and disjunct species occurring in low-
land tropical forest and with perhaps older origins.

The present collaborative project was undertaken at
the Natural History Museum, London, with the sup-
port of the Royal Society (London) and NATO (see
Puplesis & Robinson, 1999). It combines Robinson’s
interests in Microlepidoptera biodiversity with
Puplesis’s expertise and interest in inventorying the
families of primitive Microlepidoptera. We hope that

8

re-examination, characterisation and illustration of the
‘old types’ together with an overview using newly-
collected material will stimulate further studies of the
Nepticulidae in this much-neglected continent. The
current publication is intended to provide a rough
foundation upon which we and others can build a more
detailed account of the diversity of the Neotropical
Nepticulidae.

MATERIAL AND METHODS

Primary types of practically all species were exam-
ined; depositories of types are given in the
redescriptions of species. Additional material (mostly
unidentified specimens) available in the BMNH col-
lection as well as Neotropical material available from
other institutions (ZMUC, USNM and VPU) was stud-
ied. Material studied includes that collected recently
by the senior author and S.R. Hill in Belize (1998) and
by colleagues in Mexico, Dominica, Venezuela, Peru,
Chile and Argentina (1963-1987).

In Belize larvae were collected and reared, and
adults were collected at light. Mined leaves (or other
plant parts) were placed in Petri dishes which were
then checked regularly for emerged adults. Emergence
occurred within 1-2 weeks. Adult moths were col-
lected by attracting them to mercury-vapour light from
a lamp suspended slightly above eye-level and 5—
10 cm in front of a white screen, rather closer than is
usual in the standard method for light-collecting
(described by Robinson et al., 1994) in which the lamp
is about 0.5 m from the illuminated surface. A Honda
EX 350 generator was used as a power-source. As
many different habitats and sites as possible were
sampled. Moths attracted to the screen were collected
into small glass tubes and pinned after killing with
ethyl acetate.

Genitalia were prepared following the method
described by Robinson (1976). After maceration of the
abdomen in 10% KOH and subsequent cleaning, male
genital capsules were removed from the abdomen and
mounted ventral side uppermost. Where the genital
armature was particularly complicated, the genitalia
were studied and sketched in glycerin before perma-
nent mounting. The aedeagus was removed and
mounted alongside the genital armature except in the
case of some paratypes or where its removal would
jeopardise study of fine structure and where it did not
obscure other sclerites. Female genitalia were removed
entirely from the abdomen, cleaned and mounted ven-
tral side uppermost. Genitalia and abdominal pelts of
both sexes were stained with Chlorazol Black (Direct
Black 38/Azo Black) or, occasionally, mercurochrome,
and mounted in Euparal.

Forewing length is expressed as a range, where

R. PUPLESIS AND G.S. ROBINSON

availability of material made this possible, measured
along the costa from the wing base to the apex of the
cilia. Wingspan was measured from the tip of the left
wing to the tip of the right wing, where well-mounted
specimens were available; in other cases the forewing
length was doubled and the thorax width added. Anal
tufts of males have not always been described because
they do not survive on worn specimens and tend to be
lost when specimens are ‘relaxed’ prior to setting.

Illustrations of adults were drawn in Indian ink by
Mrs Birute Noreikiene (VPU) using preliminary
sketches and notes by Puplesis and with additional
observations using a stereoscopic microscope. Adults
are illustrated at the same scale to indicate comparative
size. The condition of many of the specimens studied is
poor, and the illustrations of adults are therefore ideal-
ized and generalized. In a few cases, they could be
described as reconstructions from damaged specimens.
Unfortunately black and white drawings cannot show
the metallic lustre and coloured iridescence character-
istic of most Nepticulidae; details of such colours have
been incorporated into species descriptions.

Genitalia and wing venation drawings were made
by Puplesis using a camera lucida, mainly from perma-
nent slides, but occasionally from temporary glycerin
mounts (see above).

Types of all newly described species are deposited
in the institution from which the specimen
was received, indicated in the species description.
Described but unnamed species are distinguished by
the number of the corresponding genitalia slide, e.g.,
“Acalyptris species 29135’ is a taxon exemplified by
the specimen from which male genitalia slide 29135
(BMNH collection) was made.

ABBREVIATIONS OF INSTITUTIONS

ANS Academy of Natural Sciences, Philadelphia,
Pennsylvania, USA

BMNH The Natural History Museum, London, UK
(formerly British Museum (Natural History))

FSCA Florida State Collection of Arthropods,
Gainesville, Florida, USA

MACN Museo Argentino de Ciencias Naturales, Buenos
Aires, Argentina

NNM Nationaal Natuurhistorisch Museum (Naturalis),

Leiden, Netherlands
VPU Vilnius Pedagogical University, Vilnius,
Lithuania

USNM _ National Museum of Natural History, Washing-
ton DC, USA (formerly United States
National Museum)

ZMUC Zoological Museum, University of Copenhagen,

Copenhagen, Denmark

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

ACKNOWLEDGEMENTS. Ole Karsholt and Niels P.
Kristensen (ZMUC) provided the initial stimulus for the
present project together with generous support during its
course. We thank Don R. Davis (USNM), Ole Karsholt
(ZMUC) and Erik van Nieukerken (NNM) for the loan of
material and for providing valuable information; Arunas
Diskus (VPU) for his painstaking help in much of the dissec-
tion; Birute Noreikiene (VPU) for the drawings of adults and
of the mines of Acalyptris; Caroline Whitefoord (BMNH) for
the identification of Lonchocarpus lineatus from Belize.

The first author is extremely grateful to Kevin R. Tuck
(BMNH) for generous support during the course of this
project and especially to Simon R. Hill (University of West-
minster, UK) for his selfless and generous help with planning
fieldwork, transport of equipment, technical assistance and
companionship during the Belize expedition in 1998. He also
thanks: Alex Monro and Pat Haynes (BMNH) for pre-field-
work advice; the staff of the Las Cuevas Research Station,
Belize, particularly Chris Minty, for support; the owners of
Pook’s Hill Nature Reserve, Belize, Vicky and Ray Snaddon,
for their hospitality and interest in the project; Arunas Diskus
and Virginijus Sruoga (VPU) and particularly Kasparas
Puplesis for technical assistance with preparation of equip-
ment; Jurate Puplesiene (Antwerp), for enthusiastic help with
descriptions of mainly Argentinian species; Zita Puplesiene
for providing help and support during the final preparation of
the manuscript. Special thanks are due to Paul Southwell,
Bruce Agnew and Jim and Marie Hill for their generous
hospitality and friendliness during Puplesis’s long-term stays
in Britain. Patrick and Amanda Hills are thanked for their
friendly interest in the project.

This project was conducted during tenure by Puplesis of a
one-year NATO Postdoctoral Fellowship (sponsored by NATO
and administered by the Royal Society of London) at the
Natural History Museum and finalized during a study visit to
BMNH under the Royal Society/Lithuanian Academy of
Sciences Exchange Programme in 1999. We are most grateful
to the Royal Society of London, NATO, the Godman Explora-
tion Fund (Hon. Secretary — Judith Marshall) and the Prof
Hering Memorial Research Fund (Hon. Secretary — Malcolm
Scoble) for support. Puplesis thanks the Trustees and the
Keeper of Entomology of The Natural History Museum,
London, for study facilities and access to collections. We
thank Malcolm Scoble, Klaus Sattler and Kevin Tuck for
reviewing this paper, and for their many perceptive and useful
suggestions.

MORPHOLOGY AND TAXONOMIC
COMPOSITION OF THE FAMILY

Adult morphology

The morphology of nepticulids has been extensively
discussed by, particularly, Scoble (1983), Nieukerken
(1986b), Johansson et al (1990) and Puplesis (1994).
Conspicuous morphological features are discussed
below to provide a background to understanding the

9

morphological particularities of the newly describing
taxa and justifying amendments in terminology.

The head-capsule of nepticulids is hypognathous,
approximately oval, and slightly flattened dorsoven-
trally. Only a few sulci are present on the head-capsule.
The clypeus is a small triangular or rounded lobe; a
clypeal sulcus is absent. The mouthparts normally are
only developed weakly; mandibles are absent; the
maxillae are represented by galeae and maxillary palpi;
laciniae are absent; the galeae together unite to form
the proboscis (= haustellum) which may sometimes be
functional. Occasionally the galeae are very long; the
maxillary palpi are always five-segmented, with
sensilla; the labial palpi are normally three-segmented
and widely separated at their bases, occasionally two-
segmented; the first (basal) segment of the labial palpus
is usually longer than the second and third segments. In
slide preparations of the head-capsule, the sucking
pump and membranous hypopharynx are usually vis-
ible. Ocelli are absent; chaetosemata are present. The
compound eye is generally large, its size relative to that
of head-capsule varying little. The eye-index (Powell,
1973) is fairly high, in the range 0.9-1.2, and is possi-
bly related to time of flight (diurnal species, according
to Powell (1973), have a lower index than night-flying
ones). There are no striking differences in eye-index
among the Neotropical species dealt with here.

The antenna is always shorter, usually much shorter,
than the forewing, its basal segment (scape) strongly
enlarged and forming an eye-cap with a concave un-
derside. The eye-caps cover the compound eyes when
the moth is at rest. The flagellum comprises 12 to 70
segments (flagellomeres) that are more or less uniform
except for the first (the pedicel) which is longer than
the rest. Four types of sensilla have been found on the
flagellomeres of Nepticulidae: sensilla trichodea, s.
chaetica (two types), s. coeloconica and s.
vesiculocladum (see Nieukerken & Dop, 1987). The
sensillum vesiculocladum described by these authors
occurs in all species examined but is unknown outside
the family.

The tufted head is characteristic. Piliform scales
arise from a large circular area on the front of the head,
and also from two other patches on the vertex and are
collectively named the frontal tuft; its colour has diag-
nostic value. Two groups of piliform or lamellar scales
are attached to the back of the head, often overlapping
the thorax, and collectively termed the collar. The
frons is smoothly scaled below the frontal tuft.

The colour pattern of head, thorax and forewing is a
very important diagnostic feature; however it is of
almost no phylogenetic use, since Neotropical
nepticulids (like those from other regions) show great
homoplasy. A partial exception is Manoneura, with
strikingly strong blue-purple lustre, and possibly a few
Neotropical Enteucha with unique white-tipped
forewings. The forewing may be uniform or patterned

10

(with one to three fasciae, with spot(s) or with both
fasciae and spot(s)). Most nepticulids have uniformly
grey or blackish forewings with a single fascia, the
dark areas often with bright metallic lustre which may
be bronze, purple, blue or green. But forewing patterns
are more diverse in the Neotropical species described
here: nearly 30% have a single white/light fascia on a
dark background, 20% have irrorated forewings (ir-
regularly dispersed darker scales on a paler
background), 14% have a dark fascia on a paler back-
ground, 12% are unicolorous, 6% have three fasciae on
a dark background; other species have two fasciae ona
dark background, two opposing pale spots, and one
spot. Very unusual pattern types among the Neotropical
nepticulids include Enteucha cyanochlora with a tri-
angular costal spot together with a white forewing tip,
and two species (Stigmella andina and S. pruinosa)
with striking sexual dimorphism: females of andina
have distinctive black-tipped forewings whereas in
males the forewing is uniformly pale; males of pruinosa
have a broad band of whitish androconial scales over
brown forewings, absent in females.

Generally fasciae or spot(s) vary from white to
yellowish, with a silver or gold sheen or lustre,
occasionally other colours. In females the scales usu-
ally appear more coarse, and pale markings (if present)
are more extensive than in the male. The hindwing
varies from creamy white to dark brown, and in males
is sometimes covered by black or orange androconial
scales (but these are not known in Neotropical species).
A cilia-line, formed by dark-tipped, lamellar scales,
may be present. Androconial scales may be present in
males, and normally occur on the underside of the
forewing or either surface of the hindwings. They may
entirely cover the wing, but in other cases are distrib-
uted in patches. Sometimes, long specialized
androconial scales may extend into the fringe (see
Acalyptris trifidus, Fig. 52; Stigmella pruinosa, Fig.
34; partially S. albilamina, Fig. 26). Additionally,
Nieukerken (1986b) recorded patches of velvet-tex-
tured raised scales on the hindwing underside of
European Trifurcula.

Wing-coupling in males comprises a double mecha-
nism: firstly, a composite frenulum arising on the
hindwing base hooks into the costal retinaculum, which
is comprised of hooked scales; secondly, a row of
costal bristles (pseudofrenular bristles), arising on the
humeral lobe, couples with a subdorsal retinaculum
composed of hair-scales. In females there is no frenular
bristle, and only the second mechanism is present. For
details see Nieukerken (1986b). Some species possess
pseudofrenular bristles modified into a hair-pencil (for
example, along blackish cluster in Ectoedemia species
29105, Fig. 40).

Wing venation is reduced in Nepticulidae (Figs 61—
65), although reduction is generally not as extreme as
in Opostegidae (Puplesis & Robinson, 1999). In the

R. PUPLESIS AND G.S. ROBINSON

largest nepticulid genus, Stigmella, forewing vein Sc
is short and sometimes weakly visible. R + M coa-
lesce, and there are three radial veins (R,, Ree Re.)
Cu is present as a separate vein. Veins 1A and 2A are
fused along their entire length and are well-developed
(Fig. 63). In the hindwing, Sc +R, is short, and R, and
M either usually coalesce in the basal half of the
hindwing or are semi-reduced (as in Stigmella
albilamina, Fig. 63). Cu and A are separate veins.
However, in the Neotropical Stigmella ovata (Fig. 62),
uniquely for Stigmella, R,, R,, M, and M, are pre-
served in the forewing as four separate veins, and Cu is
very long and anastomoses with the medial branch;
this striking configuration is probably a plesiomorphy.
The most complete venation pattern 1s characteristic of
Etainia (most notably in Holarctic species). The
forewing in this genus normally exhibits one subcos-
tal, five radial, two medial, one cubital and one anal
vein together with a closed cell between M and Cu.
The most reduced type of venation within the
Nepticulidae is characteristic of Neotropical Enteucha
and Manoneura (Fig. 61). However, a few southern
African Trifurcula and Etainia, and, probably, a few
Neotropical Fomoria (Fig. 64) show independent re-
duction of the venation, a feature which limits the
value of that character for phylogenetic purposes.

The legs are well-developed, frequently concolorous
with the underside of the thorax, and varying from
silvery-white to blackish. The foreleg is without spurs
or epiphysis; the midleg has a single pair of spurs, the
two pairs on the hindleg are situated at or near the
middle and at the apex of the tibia. The position of the
proximal pair of spurs is not constant.

The pregenital segments of the abdomen, including
VII in females and VIII in males, are clothed uniformly
with lamellar scales. The underside of the abdomen is
often paler than the upperside. Tergite VIII of males
and VII of females usually bears a pair of anal tufts.
Large, paired tufts of extremely long piliform scales
arise on tergites IV and VIII in members of the Asiatic
Acalyptris repeteki species-group. They arise from T-
shaped sclerotizations and completely cover the caudal
part of the abdomen. Smaller but undoubtedly ho-
mologous tufts are present in some North American
Acalyptris species (Wilkinson, 1979; Puplesis, 1984c),
but have not been discovered in Neotropical Acalyptris.
The phylogenetic significance of these tufts (more
than half the length of the abdomen) is obscure, but
they are a useful diagnostic feature at species level.
The colour of the scales on the pregenital and genital
segments is useful for separating at least two
Neotropical species (Stigmella albilamina and S.
fuscilamina) in which this is practically the only exter-
nal difference.

The male genitalia are symmetrical except for the
slightly asymmetrical aedeagus and asymmetrically
distributed cornuti. The tegumen is fused with the

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

vinculum through the vinculum arms, completing a
ring-shaped annulus. The tegumen, a simple band, is
very variable in size. Occasionally the tegumen may be
invaginated or extended posteriorly. A pseuduncus
(posterior extension of the tegumen), if present, is
situated dorsal to the uncus or coalesced with it
(Etainia); usually the pseuduncus is a broad (Fig. 174
) or narrow lobe (Fig. 178), but it is sometimes divided
into lateral lobes (Figs 157, 164, 180, 182). There is no
boundary between the pseuduncus and tegumen.

Additional rod-like apodemes arising from the
tegumen may be present in the male genitalia of
Acalyptris. In Neotropical representatives these are
still not fully separated from the tegumen (Figs 170,
172, 193) or are very under-developed (Fig. 178), an
apparently more plesiomorphic state than in the
Palaearctic species of the genus.

The uncus is narrowly V-shaped, thickened or hood-
like; it is often bilobed or even with three narrow
processes (Fig. 184) and rarely takes other forms (Figs
148, 164). The unusual uncus with a single anterior
process in Manoneura (Figs 84, 86) has been mis-
interpreted as ‘downfolded’ (Nieukerken, 1986b).
Occasionally, an uncus is absent (Ectoedemia — Figs
152, 154).

The gnathos is a small sclerite connected through its
lateral arms to the tegumen (Figs 80, 90, 148, 164,
196). It consists of a transverse bar (occasionally not
developed), posterior processes (or process) and lat-
eral arms. Sometimes anterior processes are
additionally developed, and in other cases posterior
processes can be replaced by a central element. An
unusually broadened posterior process in Acalyptris
platygnathos partially replaces the central plate (Fig.
202). A very unusual modification of the gnathos is
observed in three Neotropical species united below
into the new Stigmella eurydesma species group: the
anterior plate-like part of the sclerite is strongly re-
duced, but the posterior part is developed into very
strongly sclerotized and melanized lateral processes.
These processes partially coalesce with the more dor-
sal uncus, and it is sometimes difficult to recognise two
separate sclerites (Figs 120, 126, 129). The very unu-
sual gnathos in Manoneura (Figs 84, 86) has been
misinterpreted as a complex structure comprising
gnathos+uncus as a single fused sclerite (Davis, 1978).
Together with the tegumen, the gnathos surrounds the
tuba analis. Slightly sclerotized and sometimes spined,
the dorsal part of the tuba (very rarely visible in
Nepticulidae) is referred to as the scaphium, and its
ventral region as the subscaphium (Klots, 1956).

The valvais always strongly sclerotized and exhibits
numerous diagnostic differences between species. The
apical region usually tapers to a variable distal process,
in some species (predominantly in the Stigmella salicis
group) to two processes (e.g., Figs 96, 97). In some
taxa the apex of the valva remains broad. The inner

ill

margin of the valva may bear one or more well-
developed lobes or processes (Figs 174, 196), which
may be numerous and elaborate in some Neotropical
species (e.g., Figs 79, 157, 159, 202). In Fomoria
species 29122 the valva is actually divided into two
lobes (Fig. 161). Numerous setae are distributed on the
valva, especially on the inner margin and at the apex;
Stigmella barbata has very long, strong, hair-like setae
from the apical process (Fig. 140) and in this respect
resembles a few Nearctic species. In some cases (nota-
bly in the Australian Pectinivalva Scoble and some
Asiatic species of Acalyptris) the valva bears a
pectinifer, more or less resembling the pectinifer of
opostegids. There is still no general agreement as to the
homology of these pectinifers. However, the pectinifers
of the Asiatic Acalyptris falkovitshi (Puplesis),
Pectinivalva sp. and Opostega auritella (Hiibner) have
been well illustrated and compared by Nieukerken
(1986b: figs 60-64), and we believe they are homolo-
gous. Only a single species in the Neotropical fauna
(Acalyptris bovicorneus) has been discovered with a
similar pectinifer (Fig. 166); other related Acalyptris
species have none. The presence of a pectinifer may be
either a parallelism, or more likely a plesiomorphy
preserved in Pectinivalva and some Acalyptris; the
latter explanation gains some support from the wide-
spread but sporadic occurrence of this structure
elsewhere in the monotrysian Heteroneura, notably in
Incurvarioidea. The pectinifer in Opostegidae, sister
group to Nepticulidae, is unambiguously derived in
comparison with the hypothetical ancestral state which
appears to be expressed in those Nepticulidae with a
pectinifer. In Opostegidae the pectinifer is borne on a
cucullar lobe well separated from the main body of the
valva, a striking autapomorphy of the family expressed
in all the taxa known.

The transtilla forms a dorsal connection between
the apodemes of the valvae. The fully developed
transtilla (with a transverse bar) is reduced or absent
only in Pectinivalva, Enteucha, Glaucolepis and most
world species of Acalyptris (but in less than a third of
the Neotropical species). The Belizean Ectoedemia
fuscivittata also exhibits secondary loss (reduction) of
the transverse bar of the transtilla, not previously
recorded for the genus. In other nepticulids the transtilla
is well-developed, consisting of an uninterrupted (Fig.
164) or medially interrupted (Figs 144, 150) transverse
bar.

The term ‘sublateral process(es) of transtilla’ is
widely used in the literature on Nepticulidae and for
the most part refers to the valval apodeme. Admittedly,
it is difficult in some cases to determine where the
transtilla bar ends and the apodeme begins, but the
term is inappropriate; we use it here with considerable
misgivings. “Sublateral processes’ are invariably de-
veloped (Fig. 174), sometimes strongly so (Fig. 103)
but the fusion of the apodeme and transtilla may be

12

such that the former is indistinguishable and the whole
is a continuous rounded band of sclerotization (Figs
88, 92) that could be interpreted entirely as transtilla;
where the continuous structure is angular (Figs 104,
159) it could be inferred that the transverse element is
the transtilla and the longitudinal elements are the
apodemes, but there is no justification for this assump-
tion. Where the apodeme runs anteriorly beyond the
transtilla bar we have followed accepted practice and
designated this as a “sublateral process’, 1.e., “sublateral
process’ is asynonym of ‘apodeme’; where it does not,
we have recorded the sublateral process as absent, but
the apodeme may well still be present.

There is some intraspecific variability in the transtilla
and apodemes; where slender sublateral processes are
present, their curvature may vary; broad processes
may vary in shape and quite often the left process may
not be symmetrical with the right (Figs 103, 106).
Although the presence or absence of the transverse bar
of the transtilla is a good diagnostic feature of a few
genera, and may have some phylogenetic significance,
the characteristics of size and shape (discounting
intraspecific variability) are significant only for species
diagnosis. We treat a well-developed transtilla with a
broad and enlarged transverse bar or specialized
sublateral processes as a derived feature.

The juxta is a small triangular, trapezoidal or irregu-
larly shaped sclerotization of the membrane between
the valvae ventral to the aedeagus. It is variable in
shape and in its degree of sclerotization and occurs
irregularly in different genera of Nepticulidae with
little phylogenetic predictability. However, the juxta in
Stigmella (Fig. 97) is simpler and more plate-like, than
in, for example, Acalyptris, where it tends to be more
complicated (Figs 180, 182) or very complicated (Fig.
170) and probably contains elements derived from the
posterior extension of the vinculum. In some Stigmella
species the juxta is apparently not connected to the
vinculum; sometimes the juxtal region is entirely mem-
branous (1.e., a juxta is absent) (Fig. 96).

The vinculum is shaped like a large plate and is
sometimes narrowed or bilobed anteriorly. Two types of
vinculum are present in Nepticulidae. In the first, the
vinculum is connected through wide and well-devel-
oped lateral arms with a large tegumen and these
together form a ring surrounding the aedeagus (‘ring-
shaped’ vinculum of Beirne, 1945). In the second type
the vinculum has moderate lateral arms which connect
with a strongly reduced and usually small tegumen (“U-
shaped’ vinculum of Beirne, 1945). In the present
treatment, the complex of vinculum plus tegumen is
regarded as an annulus (following Kuznetsov, 1915).
The size of the ventral plate of the vinculum, lateral lobes
and anterior invagination may vary strongly between
species. The anterior region of the vinculum has been
referred to as the saccus (Beirne, 1945: Wilkinson &
Scoble, 1979; Newton & Wilkinson, 1982).

R. PUPLESIS AND G.S. ROBINSON

The aedeagus, in contrast to many other Lepidop-
tera, is broad and strongly sclerotized, cylindrical or
retort-shaped, frequently broadened basally and
occasionally apically; it is often 2—3 times longer than
the valva. Carinae may be developed towards the apex
of the aedeagus (e.g., Figs 153, 192, 200) and vary
considerably between species. The vesical cornuti vary
from very small and spine-like or almost granular to
large, horn-like, lamellar, or elongate and irregular.
The number of cornuti varies interspecifically from a
few to several hundred. They have great diagnostic
value, in most cases at species level, very occasionally
at higher taxonomic levels (e.g., in Trifurcula and
Glaucolepis). The striate thickening surrounding the
base of the ductus ejaculatorius (the arcuate plate
through which the ductus passes into the vesica) was
regarded as a pair of striate plates (Schoorl et al., 1985)
and was named the cathrema by Kemperman &
Wilkinson (1985). In the Australian Pectinivalva, the
cathrema is only weakly developed and this state is
plesiomorphic. In some Stigmella species a distinct
tubular membrane (the manica) surrounds the aedeagus.
This membrane is usually spinose but such a modifica-
tion has yet to be discovered among the Neotropical
species.

Females are of the monotrysian type, with a com-
mon terminal anogenital opening. The vagina is short,
widening into a vestibulum which may sometimes be
strongly folded, and almost always protrudes laterally
to form an accessory sac in Stigmella (Fig. 208). In
Ectoedemia a distinctive spiculate pouch is developed.
In Etainia, Acalyptris and Fomoria (Fig. 219) additonal
sclerotization of the vagina-vestibulum forms struc-
tures that can be described as an antrum or sclerites of
the vestibulum. The ductus spermathecae opens into
the accessory sac or into the vestibulum if an accessory
sac is absent (Fig. 207). Usually the ductus is coiled,
and occasionally it is spiculate. The bursa copulatrix is
large, usually oval, with (Fig. 208) or without
pectinations or signa. The signa are reticulate (in
Trifurculinae sensu Puplesis, 1994) (Figs 220, 221) or
in a different pattern; sometimes they are large spines
(in some species of the Palaearctic Stigmella paliurella
species-group). In some species of the Palaearctic
Stigmella ruficapitella species-group, the accessory
sac has functionally replaced the bursa copulatrix,
which is minute or indiscernible. The apophyses are
well-developed and vary interspecifically. The anal
papillae usually form a broad, flat posterior margin to
the abdomen. The ovipositor is weakly developed,
occasionally protruding.

Biology

The biologies of Neotropical nepticulids are known
only from very scanty reared material. Generally,
nepticulid eggs are laid singly, glued to the surface of

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

a leaf or another (often damaged) plant organ. The
precise oviposition site is sometimes characteristic, as
certain species lay their eggs either on the upper- or
underside of a leaf. But other species may lay on either
side of the leaf. Usually just one egg per leaf is laid, but
in some species (when population density is high) a
few or numerous mines may be found in one leaf. For
example, in the Western Palaearctic Stigmella paliurella
(on Paliurus spina-christi) and Stigmella ficulnea (on
Ficus carica), 10 to 30 mines have been noted in one
leaf (in the Caucasus and Turkmenistan). Between 400
and 500 mines of the eastern Palaearctic Ectoedemia
picturata have been noted on one (composite) leaf of
Rosa rugosa in far eastern Russia (Puplesis, 1985).
The duration of the egg stage varies greatly, from 8-11
days to 130-150 days. Although this has some correla-
tion with temperature (Kino, 1981), it relates also to
the voltinism of the species. Four or five larval instars
are typical, but occasionally — as in the subgenus
Ectoedemia (Zimmermannia) — 6-8 instars are known
(Schonherr, 1958; Nieukerken, 1985b; Nieukerken &
Johansson, 1990). After hatching, the young larvae
bore into the leaf. The majority make upper-surface
mines, but occasionally mines are under-surface ones
(e.g., in the Asiatic Stigmella flavescens). The majority
of Nepticulidae are leaf-miners, but sometimes larvae
mine buds and shoots (Etainia, and some Bohemannia
species); the summer generation of Etainia species
makes short mines in the keys (fruits) of Acer;
Ectoedemia (Zimmermannia) mine in young bark
(cambium). The Neotropical Ectoedemia species 29105
may be a bark-miner, because similar species of this
group in the Holarctic region feed only in bark. Amongst
the numerous leaf-miners, some species (Ectoedemia
populella-group) start to mine in the petiole or midrib
of a leaf and only later enter the tissues of the leaf-
blade. Old mines are sometimes completely
transparent, or at least translucent and easily visible
(e.g., the Neotropical Stigmella gossypii, Enteucha
gilvafascia). In some species, various instars make
different types of tunnels, each type with characteristic
frass deposition (the Neotropical Acalyptris species
29140 on Lonchocarpus lineatus (Leguminosae) has
uniform frass deposition throughout the whole mine,
Fig. 6). The tunnels made by Nepticulidae have char-
acteristic patterns. Mines can be generally categorized
as: linear (most species) (Figs 4-6); blotch (very rare);
a combination of gallery in the first half and blotch in
the second half (not rare); spiral (= helical) (only in
Enteucha). Bark mines are always linear. The mines of
the majority of species are characteristic and have
diagnostic value at species level. However, the shape
of the mine and the colour and distribution of the frass
may vary, depending (amongst many factors) on the
thickness of the leaf, light conditions and hostplant
species.

The larva never leaves its mine until it is full-grown.

13

However, in most investigated Glaucolepis, and some
other species, the larvae use more than one leaf
(Nieukerken & Johansson, 1990). These species mine
continuously from one leaf to another via petiole or
stem. The duration of the larval stage is usually short,
in the majority of cases a matter of a few days (in
temperate regions, as well as in Belize), but sometimes
this stage may take a few months. A prolonged larval
stage is characteristic of most Ectoedemia, which mine
from late summer to autumn, and for species
overwintering as larvae (e.g., the Palaearctic Fomoria
weaveri and Stigmella castanopsiella). The longest
larval stages are known for species of Ectoedemia
(Zimmermannia) (Nieukerken, 1985b). The season of
the larval stage varies, but larvae of most species mine
in autumn. Larvae of certain Ectoedemia species mine
in yellowed or even fallen leaves, with the conserva-
tion of characteristic “green islands’.

Except in a few species (the Palaearctic Fomoria
weaveri group, Ectoedemia agrimoniae and Trifurcula
eurema), the fully-grown larva emerges from the mine
and descends on a silk thread, usually to the ground.
Larvae may then hide in detritus where they spin a silk
cocoon. In some species the cocoon is spun on a stem
or trunk of the hostplant. The duration of the pupal
stage is short in the summer generation(s) of bivoltine
or multivoltine species, usually 10-20 days. In the
overwintering generation, fully-grown larvae do not
pupate but diapause through the long, cold period in a
prepupal stage. After diapause, the pupal stage is as
short as that of summer generations. The pupa is
protruded halfway out from the cocoon by means of its
abdominal spines, before the moth emerges
(Nieukerken ef al., 1990).

The imago has a short lifespan. Indoors, when natu-
ral conditions are mimicked, the lifespan of the moths
amounts to 2—3 days, or 5—6 days when fed with sugar
syrup. The lifespan of adults collected in the wild and
not fed is usually only one day, rarely three days.
According to Johansson et al., (1990), individuals of
several species aestivate in southern Europe. This may
also be a characteristic of some central Asiatic species.
Otherwise, only when the imago is kept at a low
temperature (about 10°C), may the lifespan (with the
moth generally in torpid condition) be lengthened to
up to 5S—6 days. The duration of the flight period of a
species is about 20-30 days, sometimes longer. Emer-
gence starts at different times in the case of different
species. Usually, the first males emerge earlier than the
first females. The moths usually fly around their
hostplants in the afternoon and evening before dusk.
Adults of most species are attracted to light, males
being collected rather more frequently than females.

Before copulation the male exhibits specific behav-
iour in the form of a ‘dance’. The male runs close to or
around the resting female, vibrates his raised wings
and turns his abdomen slightly upward. If the female

14

responds by raising her wings slightly and vibrating
them a little, the male instantly turns his abdomen
towards her, and mating takes place. The wings of both
individuals are then lowered. Duration of copulation is
at least 45 minutes in the eastern Palaearctic Stigmella
kozlovi (Puplesis, 1984a) and about 20 minutes in the
western Palaearctic Ectoedemia liebwerdella
(Schonherr, 1958).

Univoltine species are rare among the Nepticulidae.
However, within the subfamily Trifurculinae, many
species of Ectoedemia and Trifurcula and some
Fomoria and Bohemannia have only one generation
per year. In most univoltine species the larval stage is
unusually long, especially in Ectoedemia
(Zimmermannia). In this group this stage lasts from
two to six or seven months, or even nearly two years.
Bivoltine species are in the majority in the Holarctic
region. Details of the biology of tropical species are
largely unknown. A few species have been noted as
multivoltine, but we would expect a great number of
such species in the tropics. Even among the boreal
fauna, the majority of the so-called bivoltine species
might have three (or more) generations per year if
climatic conditions permit.

Nepticulidae are trophically associated with
dicotyledonous plants from the Magnoliophyta.
According to Nieukerken (1986a, 1986b), plants from
18 subclasses and numerous families are fed upon by
nepticulid larvae. Exhaustive data on the hostplant
relationships of western Palaearctic Nepticulidae have
been presented by Nieukerken (1986a) and by
Nieukerken & Johansson (1990). The majority of
nepticulid species in the Holarctic mine plants from a
few families: Rosaceae, Betulaceae, Salicaceae and
Leguminosae. Further south, more families are in-
volved (Puplesis, 1988, 1992; Nieukerken & Johansson,
1990) But for many species the hostplant is still un-
known (Puplesis et al., 1996). Two new hostplant
families (Cunoniaceae and Eucryphiaceae) were
recently recorded by Hoare (2000) for the Australian
fauna; they were considered as a part of the ancient
angiosperm flora that covered large areas of Australia,
Antarctica and South America in the late Cretaceous
and early Tertiary (see Hoare, 2000). Currently only
five nepticulid hostplant families have been recorded
in the Neotropical Region: Leguminosae
(Lonchocarpus lineatus), Compositae (Senecio
bonariensis), Onagraceae (Ludwigia major),
Malvaceae (Gossypium barbadense, occasionally
G. hirsutum) and Polygonaceae (Coccoloba uvifera,
the hostplant of two species).

Monophagy is known to occur within the family,
but by far the greatest proportion of nepticulid species
are oligophagous (Nieukerken, 1986a; Puplesis, 1994).
Broad oligophagy is especially characteristic of species
feeding on Rosaceae. Strict oligophagy is the most
common trophic type within the family and applies to

R. PUPLESIS AND G.S. ROBINSON

nearly 80% of all investigated Nepticulidae. A few
cases of disjunct oligophagy are known in the Holarctic
fauna but no actual polyphagy. There are numerous
species among Nepticulidae which can be very abun-
dant and damage the hostplant (the Neotropical
Stigmella gossypii on Gossypium may fall in this cat-
egory). However most species are no more than
potential pests, and significant host damage appears to
be an exception brought about by particularly favour-
able ecological conditions. Nepticulid populations
appear to be able to react with very rapid increases in
density.

Taxonomic composition

The taxonomic history and classification of the
Nepticulidae has been extensively discussed by
Puplesis (1994) who proposed a cladogram based on
79 characters. In this cladogram, the Nepticulidae have
two main lineages: Nepticulinae (including the Aus-
tralian Pectinivalva) and Trifurculinae. Despite the
exotic appearance of Pectinivalva (mainly due to a
series of striking plesiomorphies), its apomorphic
features were found insufficiently unambiguous to
support its recognition as a separate subfamily. The
Nepticulinae and Trifurculinae differ mainly in the
basal structure of the annulus (see Morphology) and in
a set of correlated apomorphic genital characters of
genitalia believed to be independently derived. Within
the Trifurculinae, Bohemannia takes the lowest
phylogenetic branch, i.e., it is the sister group of the
remaining Trifurculinae (for details, see Puplesis,
1984b, 1992, 1994). However, an alternative classi-
fication proposed by Scoble (1983) is better supported
by characters of the immature stages and has often
been followed in recent literature with some adjust-
ments at generic level (Nieukerken, 1986b; Nieukerken
et al., 1990; Hoare et al., 1997; Hoare, 2000). This
latter classification recognizes two alternative sub-
families: Pectinivalvinae (including two endemic
Australian genera Pectinivalva and Roscidotoga) and
Nepticulinae (including all remaining genera divided
between two tribes, Nepticulini and Trifurculini). In
Trifurculini three genera (Fomoria, Laqueus and
Etainia) have recently been synonymized with
Ectoedemia and two (Glaucolepis and Levarchama)
are now treated as subgenera of Trifurcula (Nieukerken,
1986b; Nieukerken et al., 1990).

There is still instability in the taxonomic rank and
placement of several nepticulid genera and subgenera.
In total, we currently recognise a total of 16 genera
worldwide. Seven of these are represented in the
Neotropical region. We tend here to follow the classi-
fication of Puplesis (1994), fully cognisant of its
provisional nature. Classifications sometimes collapse
when challenged by the addition of large numbers of
tropical species, and this may be about to happen with

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

the Nepticulidae. The addition of further data from
tropical faunas may soon make necessary a reappraisal
of nepticulid generic phylogeny.

A brief discussion of each genus follows.

Synopsis of genera

Pectinivalva Scoble, 1983. Type species:
Pectinivalva commoni Scoble, 1983. This genus pre-
serves many striking plesiomorphic features such as a
pectinifer on the valva, and rounded or unspecialized
valve apex; some of these are shared with Roscidotoga:
2- or 3-segmented larval antenna, long frontal setae on
pupa (Hoare, 2000), vein 1A+2A of forewing not
thickened, weakly developed cathrema. However, at
least five autapomorphies support the monophyly of
the genus (Hoare, 2000). There are several
synapomorphies with Roscidotoga (Rs+M trunk de-
flected towards costa in hindwing; row of spines on
anterior sclerite of second sternite; loss of fourth larval
mesothoracic seta ventral to SV 1), and possibly pupal
characters (Scoble, 1983; Hoare, 2000). However, the
polarity and consistency of these synapomorphies
(which support the monophyly of Pectinivalva +
Roscidotoga and thus of the Pectinivalvinae) are per-
haps questionable. In the revised diagnosis of
Pectinivalvinae suggested by Hoare (2000), the strong-
est autapomophy of Pectinivalvinae (Rs+M trunk
deflected towards costa in hindwing) is by no means an
obvious character state (see Pectinivalva
warburtonensis (Wilson) in Hoare et al., 1997: fig.
36).

Roscidotoga Hoare, 2000. Type species:
Roscidotoga callicomae Hoare, 2000. Sister-group to
Pectinivalva; recently described from Australia, and
well-founded on 9 autapomorphic features (Hoare,
2000). Three strikingly coloured species without a
pectinifer on the valva are currently known (Hoare,
2000).

Simplimorpha Scoble, 1983. Type species: Stigmella
lanceifoliella Vari, 1955. This genus currently com-
prises just two species and has a Mediterranean —
South African distribution. It has not been recorded
from the Neotropics. The monophyly of the genus is
supported by at least three distinct autapomorphies:
Anacardiaceae as hostplant family, strongly reduced
gnathos and reduced uncus. Reduction or loss of
gnathos is parallelled in the Australian Roscidotoga
(and to a very minor extent in the Neotropical Enteucha
terricula); reduction or loss of the uncus is parallelled
in some Ectoedemia species.

Enteucha Meyrick, 1915 (= Johanssonia Borkowski,
1972; Artaversala Davis, 1978). Type species:
Enteucha cyanochlora Meyrick, 1915. This small ge-

115)

nus is probably widely distributed; it was originally
described from the Neotropics (Guyana) and, prior to
this review, comprised three described species, two
Neotropical and one European. The current study adds
four more Neotropical species so providing a broader
view of the morphology of the taxon. Enteucha may be
easily recognized by the strongly reduced wing vena-
tion, the absence of a transverse transtilla bar, the
under-developed cornuti, very large vinculum plate (in
most species) and the caudally rounded or narrowed
uncus. If the lack of development of the cornuti/cari-
nae and transverse transtilla bar (parallelled in
Pectinivalva, Manoneura, Glaucolepis, some
Acalyptris) are eventually shown to be plesiomorphies,
then Polygonaceae as a hostplant, the reduction of
venation, and usually smooth elongated cathrema may
still serve as evidence for monophyly. Manoneura,
treated as a synonym of Enteucha by Nieukerken
(1986b) and Puplesis (1994), is treated below as a
distinct genus. Short modified anterior apophyses in
the female genitalia (a synapomorphy shared with
Manoneura), a well-developed paired collar consist-
ing of lamellar scales (a synapomorphy with
Manoneura and Stigmella) together with other
characters indicate a close relationship between the
three genera.

Manoneura Davis, 1979 (=Oligoneura Davis, 1978,
preoccupied). Type species: Oligoneura basidactyla
Davis, 1978. A small, and until recently monobasic
genus which now comprises two species, Manoneura
is endemic to the Neotropics. The unusual gnathos in
Manoneura (Figs 84, 86) was misinterpreted as a
complex structure comprising the gnathos and uncus
fused to form a single sclerite (Davis, 1978). It was
further misinterpreted as ‘downfolded’ (Nieukerken,
1986b). The genus can be characterized by: 1. Well-
developed paired collar consisting of lamellar scales
(synapomorphy with Stigmella and Enteucha). 2. Scales
with very strong blue and purple lustre (homoplasy
with some Bohemannia and some Stigmella; however,
the lustre in Manoneura is more developed and more
strongly blue). 3. Strongly reduced forewing venation
(Fig. 61) (autapomorphy, but parallelled with
Enteucha). 4. Uncus with large anterior process almost
touching posterior incision of gnathos (autapomorphy).
5. Gnathos with membranous lateral arms and poste-
rior thickening, possessing a posterior incision
(autapomorphy). 6. Transverse bar of transtilla lacking
(probably a plesiomorphy, parallelled in Pectinivalva,
Enteucha, Glaucolepis and some Acalyptris). 7.
Transtilla with very long sublateral processes
(apomorphy, but frequently parallelled in other
nepticulids). 8. Vinculum very large (plesiomorphy,
frequently parallelled in Enteucha, Trifurcula and some
other nepticulids). 9. Carinae on aedeagus (apomorphy,
but frequently parallelled in other nepticulids). 10.

16

Cornuti absent (probably a symplesiomorphy shared
with Enteucha and parallelled in a few other
nepticulids). 11. Short modified anterior apophyses in
female genitalia (synapomorphy shared with
Enteucha).

Areticulata Scoble, 1983. Type species: Areticulata
leucosideae Scoble, 1983. A monobasic genus
described from South Africa, of uncertain taxonomic
position. Despite the unusual structure of the vinculum
(not re-examined by the present authors), itis probably
related to Stigmella. The genitalia possess a gnathos
with a long transverse bar and two lateral processes
directed caudally (parallelled in some Stigmella). Fora
detailed diagnosis, see the original description.

Stigmella Schrank, 1802 (=Nepticula Heyden, 1843;
Astigmella Puplesis, 1984). Type species: Phalaena
(Tinea) anomalella Goeze, 1783. This is the biggest
nepticulid genus, with a world-wide distribution and
well-represented in the Neotropical Region. It is char-
acterized by a few, generally apomorphic characters:
paired collar of lamellar scales (synapomorphy with
Enteucha and Manoneura); generally bilobed (with
possible further elaborations) uncus; gnathos with two
processes (very occasionally with a single one); large
or moderately large accessory sac in female genitalia.
Stigmella comprises many monophyletic entities which
are usually designated as species groups; the salicis
group is recognized in this study as very well repre-
sented in the Andean fauna.

Bohemannia Stainton, 1859 (= Scoliaula Meyrick,
1859). Type species: Nepticula quadrimaculella
Boheman, 1853. This genus is currently known only
from within the Palaearctic Region, and comprises 8
described species. For the intriguing peculiarities of its
morphology see Nieukerken (1986b) and Puplesis
(1994: 34, apomorphies 43-47).

Ectoedemia Busck, 1907 (= Dechtiria Beirne,
1945). Type species: Ectoedemia populella Busck,
1907. A very large, probably world-wide genus repre-
sented in the Neotropics by both subgenera: Ectoedemia
and Zimmermannia Hering. The genus is character-
ized by astriking autapomorphy — the complete absence
of an uncus (parallelled in Simplimorpha). The
neotropical E. fuscivittata is unusual in that the transtilla
is without a transverse bar (a secondary reduction).

Fomoria Beirne, 1945 (= Laqueus Scoble,
1983). Type species: Nepticula weaveri Stainton,
1855. Large, probably worldwide genus (but not
recorded from the Australian Region). Related to
Ectoedemia. Both have the same ground plan of wing
venation, but in Fomoria the tendency towards reduc-
tion is greater: vein Cu in the forewing tends be

R. PUPLESIS AND G.S. ROBINSON

shortened or completely lacking; sometimes venation
is reduced to the extent that the otherwise characteris-
tic closed cell (an independent reduction parallelled in
Acalyptris and Bohemannia) is lost. The genitalia of
Fomoria and Ectoedemia follow the same ground plan,
but the uncus (which is entirely reduced in Ectoedemia)
is fully preserved in Fomoria (plesiomorphy). Although
some species of Fomoria may be grouped in mono-
phyletic units (the largest of which is perhaps the
subgenus Laqueus with 33 named species), the taxo-
nomic status of the genus is questionable, because its
monophyly is unproven; Nieukerken (1986b) has sug-
gested that it may be paraphyletic.

Acalyptris Meyrick, 1921 (= Microcalyptris Braun,
1925; Weberia Miiller-Rutz, 1934; Niepeltia Strand,
1934; Weberina Miiller-Rutz, 1934). Type species:
Acalyptris psammophricta Meyrick, 1921. World-wide
genus, well represented in Central America, with un-
ambiguous taxonomic status. However, there are almost
no apomorphies shared by all or almost all species. The
genus is characterized best by: 1. Closed cell in forewing
shifted towards wing-base (the strongest autapomorphy
of the group, but not always easily recognized in
individual specimens). 2. Vein Cu reduced in forewing
(apomorphy parallelled in some other nepticulids, not
occurring in all Acalyptris). 3. Forewing pattern
coarsely irrorated or with patches or dark fasciae formed
from scales similar to those that form the irroration
(not present in all species; similar patterns occur in
other nepticulid genera). 4. Large, paired tufts of
extremely long piliform scales arise on abdominal
tergites IV and VIII (unique feature among nepticulids,
distributed only among some Asiatic and Nearctic
Acalyptris). 5. Transverse bar of transtilla absent (in
most species, but not in the newly described Neotropical
taxa, below). 6. Lateral rod-like apodemes developed,
arising from inner posterior margin of tegumen and
running anteriorly parallel to the margin beneath the
valvae (in many, but not all, Acalyptris species);
apomorphy; in contrast to Palaearctic species, these
apodemes are still not fully separated from the tegumen
in Neotropical taxa, and the latter appear to express a
more plesiomorphous state of this character. 7. Valva
with apical pectinifer (probably a plesiomorphy, paral-
lelled in most Pectinivalva and shared by only a few
Acalyptris — the repeteki species group and a single
Neotropical species). 8. Posterior extension of ventral
plate of vinculum forming a juxta (apomorphy shared
by most species). 9. Aedeagus with well-developed
carinae (synapomorphy shared by almost all
Trifurculinae). 10. Female genitalia with rod-like or
similarly elaborated sclerites in vestibulum. This ge-
nus comprises a few identifiable monophyletic
subgroups — three of these have been recognized as
species-groups (Puplesis, 1984c, 1989, 1990, 1994;
Puplesis & Diskus, 1995).

CENTRAL AND,SOUTH AMERICAN NEPTICULIDAE

Parafomoria Nieukerken, 1983 (=Parafomoria
Borkowski, 1975 [unavailable name]). Type species:
Nepticula helianthemella Herrich-Schaffer, 1860.
Small, western Palaearctic (Mediterranean-centred)
genus recently revised by Nieukerken (1983, 1985a);
characterized by reduction of R,, ,, expansion of lateral
arms of vinculum, some reduction of corpus bursae,
loss of signa and with Cistaceae as hostplant family.

Trifurcula Zeller, 1848 (= Levarchama Beirne,
1945). Type species: Trifurcula pallidella Zeller,
1848. Known from Europe and the Mediterranean,
also South Africa (Scoble, 1980a); characterized by
forewing venation (if not reduced) with connection
between R,,, and R,, . lost, vinculum large and usually
rounded anteriorly, uncus partially bifurcate or com-
pletely divided dorsoventrally, aedeagus with
well-developed carinae and cornuti including a dis-
tinct group of needle-like or curved cornuti at one side
(Scoble, 1980a; Nieukerken, 1986b; Nieukerken &
Johansson, 1990).

Glaucolepis Braun, 1917 (= Fedalmia Beirne,
1945). Type species: Nepticula saccharella Braun,
1912. This genus was originally described with just
one Nearctic species, but numerous Palaearctic species
have been added subsequently, particularly from the
Mediterranean region. Resembling and related to
Trifurcula, Glaucolepis is best characterized by the
lack of a transverse bar to the transtilla (independently
parallelled in Enteucha, Manoneura and Acalyptris,
but not in the related Trifurcula), and a very long
cornutus lying longitudinally within the vesica
(autapomorphy).

Etainia Beirne, 1945 (= Obrussa Braun, 1915
[preoccupied]). Type species: Lyonetia sericopeza
Zeller, 1839. Etainia comprises 16 species: four from
Europe, one from the Caucasus, four from East Asia,
two from North America (including one species with a
Holarctic distribution), and four from South Africa; it
has not been recorded from the Neotropics. For a full
checklist and discussion of phylogeny within the ge-
nus see Puplesis & Diskus (1996a). Characterized by:
1.Vinculum with U-shaped posterior invagination
(autapomorphy). 2. Vesica with H-shaped sclerotization
(autapomorphy). 3. Ductus bursae with group of spines
(autapomorphy). 4. Larvae mining in buds, shoots and
fruits, and not in leaves. 5.Valva of almost all species
with large basal apodeme(s) (autapomorphy). 6. Mus-
cle m, in male genitalia attached to lateral arms of
transtilla, and not to inner side of valva as in other
Nepticulidae (Puplesis & Kozlov, 1988)
(autapomorphy). This group was treated as a subgenus
of Ectoedemia by Nieukerken (1986b) but there ap-
pears to be little justification for this. We treat Etainia
here as a distinct genus.

17

Varius Scoble, 1983. Type species: Stigmella
ochnicola Vari, 1955. The status of this monobasic
South African genus is uncertain. The forewing vena-
tion is more complete than in its close relative Enteucha
(R,,, preserved, R, and R, still separate — clearly
plesiomorphies), but it is probable that this species is
no more than a plesiomorphous member of Enteucha.
Varius is unlikely to survive as a distinct genus.

CHECKLIST OF NEOTROPICAL
SPECIES

The 29 species recorded from Belize are marked with
an asterisk (*) and fall into seven genera: Acalyptris
(14 species), Stigmella (5), Enteucha and Fomoria
(three each), Ectoedemia (two), Manoneura and
Glaucolepis (one each).

NEPTICULIDAE Stainton, 1854: 295
(=STIGMELLIDAE Hampson, 1918: 387)
Type genus: Stigmella Schrank, 1802

ENTEUCHA Meyrick, 1915
1. cyanochlora Meyrick, 1915
2. gilvafascia (Davis, 1978)
3. hilli sp. n. *
4. contracolorea sp. n. *
5. terricula sp. n.
6. snaddoni sp. n. *

MANONEURA Davis, 1979, gen. rev.
7. basidactyla (Davis, 1978) comb. n. *
8. trinaria sp. n.

STIGMELLA Schrank, 1802
The salicis group

9. andina (Meyrick, 1915)
10. cuprata (Meyrick, 1915)
11. johannis (Zeller, 1877)
12. rudis sp. n.
13. marmorea sp. n.
14. peruanica sp. n.
15. epicosma (Meyrick, 1915)
16. schoorli sp. n.
17. hamata sp. n.
18. imperatoria sp. n.
19. olyritis (Meyrick, 1915)

The eurydesma group
20. eurydesma (Meyrick, 1915)
21. albilamina sp. n. *
22. fuscilamina sp. n. *

Unattributed to a group
23. gossypii (Forbes & Leonard, 1930)
24. kimae sp. n. *

18

25. plumosetaeella Newton & Wilkinson, 1982
26. barbata sp. n. *

27. pruinosa sp. n. *

28. ovata sp. n.

29. hylomaga (Meyrick, 1931)

30. costalimai (Bourquin, 1962)

31. guittonae (Bourquin, 1962)

ECTOEDEMIA Busck, 1907
32. reneella Wilkinson, 1981
33. helenella Wilkinson, 1981
34. mesoloba Davis, 1978

35. species 29105 *

36. fuscivittata sp. n. *

FOMORIA Beirne, 1945

37. molybditis (Zeller, 1877) comb. n.
38. diskusi sp. n. *

39. species 29122 *

40. latipennata sp. n. *

ACALYPTRIS Meyrick, 1921
41. bovicorneus sp. n. *

42. martinheringi sp. n. *

43. fortis sp. n. *

44. hispidus sp. n. *

45. novenarius sp. n. *

46. lascuevella sp. n. *

47. bifidus sp. n. *

48. trifidus sp. n. *

49. tenuijuxtus (Davis, 1978) comb. n.
50. unicornis sp. n. *

51. laxibasis sp. n. *

52. bicornutus (Davis, 1978)
53. species 29135 *

54. dividua sp. n. *

55. platygnathos sp. n. *

56. species 29140 *

GLAUCOLEPIS Braun, 1917
57. aerifica (Meyrick, 1915) comb. n.
58. argentosa sp. n. *

Note. A further eight species with distribution ranges
that run from northern Florida into the northern states
of the USA and into Canada have not been included in
our revision. They are: Stigmellanigriverticella (Cham-
bers), S. castaneaefolia (Chambers), S. ostryaefoliella
(Clemens), S. myricafoliella (Busck), Ectoedemia
clemensella (Chambers), E. similella (Braun),
E. virgulae (Braun), E. obrutella (Zeller). These are
species with predominantly boreal distributions and
hostplants which have no affinities with the Neotropical
region.

R. PUPLESIS AND G.S. ROBINSON

SPECIES REVIEW

ENTEUCHA Meyrick

1. Enteucha cyanochlora Meyrick, 1915
(Figs 7, 66-68)

Enteucha cyanochlora Meyrick, 1915: 241.
Enteucha cyanochlora Meyrick; Davis, 1984: 18;
Nieukerken, 1986b: figs 87-89.

MALE (Fig.7). Forewing length: 2.8 mm. Wingspan:
about 6.0 mm. Head: palpi, frontal tuft and eye-caps
unicolorous, pale creamish ochre; collar greyish brown,
same colour as tegulae; antenna pale ochre, ca. 34
segments. Thorax grey-brown. Forewing dark brown
with intensive emerald and purple lustre; oblique post-
median costal spot triangular, cream, distinctive.
Terminal cilia yellowish cream, distinctly contrasting
with ground colour of forewing and with a few yellow-
ish cream scales forming an apical spot; cilia on tornus
brown. Underside of forewing brown. Hindwing dark
brown, not contrasting with ground colour of forewing,
cilia intense brown. No androconia on forewing or
hindwing. Legs pale ochre with some grey. Colour of
abdomen unknown.

FEMALE. Unknown.

GENITALIA 6 (Figs 66-68). Capsule 390-400 um
long. Uncus band-shaped, not extended caudally, so
indistinct. Tegumen simple, short, not extending into a
pseuduncus. Gnathos with large, anteriorly rounded
central plate, small oblique caudal process and long
lateral arms. Valva 241—245 um long, triangular, pointed
apically, longitudinally divided with long dorsal lobe.
Transtilla absent (i.e., no transverse bar); sublateral
processes extending from bases of valvae relatively
long and slender. Vinculum long, anteriorly rounded.
Aedeagus 354 um long, slender. Vesica with group of
tiny cornuti distributed beyond middle, a spinose
sclerotization at apex and another smaller sclerite just
before apical spine.

BIOLOGY. Adults collected in February.

DIAGNOSIS. Externally easily distinguished (within
the genus and amongst other Neotropical nepticulids)
by the characteristic, triangular costal spot on the
forewing and by the very characteristic and unique
gnathos (Fig. 66).

DISTRIBUTION. Guyana.

CONDITION OF TYPE MATERIAL.
well-preserved.

Relatively

MATERIAL EXAMINED.
Holotype 6, Guyana [British Guiana]: Bartica, 11.1913
(Parish), genitalia slide no. 19273 (BMNH).

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

REMARKS. This is the type species of the genus.
However, recent discoveries of other Enteucha have
shown that the gnathos in this species is not typical of
the genus as a whole. Other distinctive features, such
as the divided valva, spines on the apex of the aedeagus,
the compact group of tiny cornuti, and the narrow
band-shaped uncus suggest that cyanochlora occupies
a somewhat isolated phylogenetic position within
Enteucha.

2. Enteucha gilvafascia (Davis, 1978)

Artaversala gilvafascia Davis, 1978: 221-223.
Enteucha gilvafascia (Davis); Nieukerken, 1986b: 49—
54, figs 90-92.

MALE. Described and figured by Davis (1978: 221—
223, fig. 5), that description summarized here. Forewing
length: 1.6-1.8 mm. Head entirely and uniformly
stramineous, almost white; antenna fuscous dorsally,
whitish ventrally. Thorax fuscous. Forewing fuscous
with a broad sharply defined pale yellowish to nearly
white median fascia and a similarly coloured wing
apex (before pale stramineous cilia). Hindwing pale
grey.

FEMALE. Similar to male.

GENITALIA 6. Described and figured by (Davis, 1978:
223, figs 29-31) and figured by Nieukerken (1986b:
figs 90-92). Uncus large, with broadly rounded poste-
rior lobe. Tegumen simple, short, not extended into a
pseuduncus. Gnathos very unusual, a transverse bar
with corners extended both posteriorly and anteriorly.
Valva divided into slender ventral lobe (cf. gilvafascia)
and broad dorsal lobe that is bent slightly inward in
apical one-third to one-quarter. Transtilla absent (i.e.,
no transverse bar); sublateral processes extending from
bases of valvae moderately short (half of width of
uncus). Vinculum very large, its extended ventral plate
distinctly triangular. Aedeagus relatively broad and
short (equal in length to vinculum); no cornuti or
apical spines.

GENITALIA 2. Described and figured by Davis (1978:
223, fig. 34).

BIOLOGY. Hostplant: Coccoloba uvifera (L.)
(Polygonaceae). According Davis (1978), the mine is
extremely long and narrow (Davis, 1978: fig. 36) and
more visible on the upperside of the leaf. Most fre-
quently the mines occur near the margin of the leaf and
only occasionally cross the main midrib. Cocoon whit-
ish to pale greyish, ca. 2-2.3 mm in length, frequently
attached to the leaf surface or other debris. As far is
known, adults fly in April to late June and early Octo-
ber to early January.

DIAGNOSIS. Differs distinctively from other Enteucha

19

and other Neotropical nepticulids in the shape of
transverse band-like gnathos, the triangular vinculum
and very deeply divided valva (cyanochlora also has a
divided valva, but the dorsal lobe is underdeveloped in
comparison with the ventral one; in gilvafascia the
opposite is the case). From snaddoni it differs in the
unreduced tegumen, very deeply divided valva, and
apically broad dorsal lobe of the valva. See Remarks.

DISTRIBUTION. Currently known from its type local-
ity, coastal southern Florida (USA), only. This species
appears to be a natural element of the Neotropical
fauna.

REMARKS. Like cyanochlora, this is a species which
by nepticulid standards is positively exotic and is a
very distinctive Neotropical exemplar of this family. It
is closely related to snaddoni, which we consider to be
its sister-group.

3. Enteucha hilli sp. n.
(Figs 8, 69, 70)

MALE (Fig. 8). Forewing length: 3.3-3.4 mm. Wing-
span: about 7.4mm. Head: palpi brownish cream;
frontal tuft pale orange-ochre; collar cream, distinc-
tive, consisting of broad lamellar scales, practically
unpaired; eye-caps cream, large; antenna brownish,
consisting of 40 segments. Thorax, tegulae and
forewing grey-brown, coarsely but indistinctly irrorated
with brown scales. Cilia brown to brownish. Underside
of forewing brown or brownish. Hindwing lanceolate
but tending to be rather wide; grey brown, almost
concolorous with forewing, cilia brown to brownish.
No androconia on forewing or hindwing. Legs brown-
ish cream, darkened with fuscous brown on anterior
surfaces. Colour of abdomen unknown.

FEMALE. Unknown.

GENITALIA 6 (Figs 69, 70). Capsule 540-545 um.
Tegumen band-like laterally, slightly extended into
papillate sublateral lobes. Uncus rounded with single
(not paired) well-sclerotized caudal papilla. Gnathos
with large posterior process and almost equally devel-
oped lateral arms; central plate weakly developed, a
narrow band; anterior processes absent but a pair of
small distinct anterior papillae present. Valva 310-
325 pum; apical one-third constricted into a very long,
slightly inward-curved process; basal two-thirds very
broad, with long distinct caudal papillae. Transtilla
absent (i.e., no sclerotized transverse bar), but basal
region of valva with long sublateral processes. Vincu-
lum long, triangular, smoothly rounded anteriorly; no
anterior emargination or lateral lobes. Aedeagus 432—
445 um, without any visible cornutus on vesica, but
with a large rhomboidal sclerite (possibly a cathrema)
basally.

20
BIOLOGY. Adults collected in April.

DIAGNOSIS. Differs from all other Enteucha by the
uniformly coloured forewing, and combination of tri-
angular vinculum with rounded uncus and basally very
broad valva. Somewhat similar and probably closely
related to the European acetosae and Neotropical
contracolorea, but separable by the characters above
which serve also to distinguish hilli from Neotropical
nepticulids in other genera.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The single specimen
is in fair condition, the wings are not spread and
slightly rubbed.

MATERIAL EXAMINED.

Holotype 3, Belize: Chiquibul Forest Reserve, Las
Cuevas, 3—16.iv.1998 (Puplesis & Hill), genitalia slide
no. 28967 (BMNH).

REMARKS. This remarkable species is named in hon-
our of Mr Simon Richard Hill, thanks to whose devoted
and enthusiastic efforts many new species (including
this one) were collected in Belize in 1998. It shares
apomorphies with all Palaearctic Enteucha (the Euro-
pean acetosae and unnamed Japanese species figured
by Nieukerken (1986b: figs 96—100)), 1.e., a single-
process gnathos, basally broadened valva and rounded
uncus. As the closest relative of the Palaearctic
Enteucha, its discovery begs several questions regard-
ing the phylogenetic and biogeographic origins of this
genus.

4. Enteucha contracolorea sp. n.
(Figs 9, 71-74)

MALE (Fig.9). Forewing length: 2.7—2.8 mm. Wing-
span: 6.0—6.2 mm. Head: palpi cream; frontal tuft pale
yellowish cream; collar comprised of dark brown la-
mellar scales; eye-caps pale yellowish cream, large
and elongated (longer than is usual in most nepticulids);
antenna pale yellowish cream (at certain angles app-
earing ochreous cream), with ~34 segments. Thorax,
tegulae and forewing smooth-scaled, brown, with gold
and weak bluish lustre. Cilia distinctly contrasting
with forewing, pale yellowish cream (concolorous with
head), brown at tornus. Underside of forewing brown.
Hindwing lanceolate, brownish; cilia brownish. No
androconia on hindwing or forewing. Legs yellowish
cream, laterally blackish brown. Abdomen dark brown
on upperside, with additional cream scales on under-
side; short rather indistinct anal tufts composed of
cream scales; anal segments, visible from underside
only, brownish cream.

FEMALE. Unknown.

GENITALIA 6 (Figs 71-74). Capsule 365-390 um.

R. PUPLESIS AND G.S. ROBINSON

Tegumen extended into short sublateral papillate lobes.
Uncus distinctive, inverted ‘V’-shaped, with small
papilla-like caudal process. Gnathos with large
posteriorly rounded caudal process, short but wide;
each arm with anteriorly-directed papilla on ‘elbow’
where the arm is reflexed caudally; central plate of
gnathos not developed, i.e., absent. Valva 195-219 um,
very broad [the preparation — 29114 — from which
Fig. 71 was drawn is slightly squashed and the valvae
look unnaturally broad]; distally constricted into huge
apical process, with spine-like process on inner basal
corner. Juxta absent. Vinculum very large, ventral plate
narrowing anteriorly, broadly rounded at apex, without
lateral lobes or anterior emargination. Aedeagus 340—
358 um, a long and relatively narrow tube, without
distinct cornuti or carinae at apex (Fig. 74).

BIOLOGY. Adults collected in April.

DIAGNOSIS. This species may be separated from all
other Enteucha by the distinctive inverted ‘V’-shaped
uncus (see also Remarks) which feature, together with
the basally broad valva and lack of a transtilla, also
separates it from Neotropical Nepticulidae in other
genera.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The single specimen
available is well-preserved, with clear scaling; the
hindwings are detached and stored in a capsule pinned
beneath the specimen.

MATERIAL EXAMINED.

Holotype d, Belize: Chiquibul Forest Reserve, Las
Cuevas, 3—16.iv.1998 (Puplesis & Hill), genitalia slide
no. 29114 (BMNH).

REMARKS. This species may be most closely related
to hilli, but it is distinctive and somewhat isolated in
possessing a gnathos of an autapomorphic type that is
quite unlike that seen in any other species of Enteucha.

5. Enteucha terricula sp. n.
(Figs 10, 75-79)

MALE (Fig.75). Forewing length: 2.7-3.3 mm. Wing-
span: 6.1—7.4 mm. Head: palpi grey-black; frontal tuft
ferruginous; collar relatively large, comprising two
groups of fuscous lamellar scales with purple and gold
lustre; eye-caps fuscous, almost black, with purple
lustre; antenna fuscous except for whitish apex, ca. 63
segments. Thorax, tegulae and forewing fuscous brown
with strong purple and some gold lustre. Medial fascia
of forewing almost straight, glossy pale gold; cilia
fuscous. Underside of forewing fuscous. Hindwing
relatively broad, grey-brown with purple lustre; cilia
brown. No androconia on hindwing or forewing. Legs
and abdomen fuscous; anal tufts of abdomen com-

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

prised of fuscous grey scales; genital segments dark
grey in ventral view.

Unknown.

GENITALIA 6 (Figs 75-79). Capsule about 600 um.
Tegumen extended into a roughly papillated and dis-
tinct lobe which analogous to pseuduncus (Fig. 77).
Uncus broad, gradually narrowed towards rounded
apex. Gnathos (Fig. 78) with broad but partly reduced
caudal process; central plate and lateral arms of gnathos
moderately developed but strongly sclerotized. Valva
320-328 pm, broad with very long apical process and
highly elaborated trilobed inner surface (Fig. 79).
Transtilla absent (i.e., no transverse bar), but sublateral
processes (apodemes) of valvae broadened basally and
very close to each other, spatulate anteriorly. Juxta
absent. Vinculum very large, ventral plate gradually
narrowed anteriorly, but broadly rounded at the end; no
lateral lobes or anterior emargination. Aedeagus rela-
tively small, 300-310 um, slightly broadened medially,
with elongate ventral sclerotization and some weakly
sclerotized spine-like cornuti; base of aedeagus tube
strongly sclerotized but not rounded as in most
nepticulids; no carinae at apex.

FEMALE.

BIOLOGY. Adults collected in late March.

DIAGNOSIS. Differs strikingly from all other Enteucha
(and all other nepticulids) in practically all major
features of the genitalia: partially reduced gnathos,
oar-like sublateral processes, multilobed valva; the
large lamellar fuscous collar and fuscous eye-caps are
also distinctive but not uniquely so. See Remarks.

DISTRIBUTION. Peru; specimens were collected at
3000 m.

CONDITION OF TYPE MATERIAL. Fair.

MATERIAL EXAMINED.
Holotype 3, Peru: Department Puno, 5 km E Limbani,
28.111.1987 (Karsholt), genitalia slide no. Diskus 183
(ZMUC).

Paratype d, data and depository as holotype.

REMARKS. This is a remarkable species — a
Neotropical ‘exotic’ — with unique multilobed valva
with oar-like processes, reduced gnathos, huge vincu-
lum, and very small aedeagus with long sclerotization.
These autapomorphic features define terricula as very
isolated within Enteucha. However, it still fits well
within the generic concept and we have no misgivings
about its placement. The adjacent broadened bases of
the sublateral valval processes are an interesting fea-
ture and may represent the development of a structure
analogous to the transtilla. The partial reduction of the
caudal process of the gnathos is a rare case of reduction
of a sclerite which is usually very conservative and
little modified in this family. Valval modifications are
widely observed, however; in terricula they are re-

21

markable and may be functionally connected with the
tiny aedeagus, providing more precise juxtaposition
than is usually required during copulation.

6. Enteucha snaddoni sp. n.
(Figs 11, 80-82)

MALE. Forewing length: ca. 2.0mm. Wingspan:
ca. 4.5 mm. Head: palpi cream; frontal tuft pale ochre;
collar cream, comprising two tufts of lamellar scales;
eye-caps cream; antenna brown on upperside and
almost cream on underside, ca. 24 segments. Thorax
and tegulae dark brown. Forewing dark brown with
wide cream medial fascia; apex of forewing also cream,
contrasting with ground colour of forewing and
concolorous with cream outer area of cilia; tornal area
of cilia brown. Underside of forewing brownish.
Hindwing lanceolate, brownish, cilia brownish. No
androconia on forewing or hindwing. Legs yellowish
cream shaded with dark brown or fuscous. Abdomen
brown on upperside and underside, small; genital seg-
ments indistinct, cream.

FEMALE (Fig. 11). Similar to male. Forewing length:
ca.2.3 mm. Wingspan: ca.5.2 mm.

GENITALIA 6 (Figs 80-81). Capsule 346-368 um
long. Uncus a large oval lobe, rather weakly sclerotized
(Fig. 81).Tegumen partially reduced (Fig. 81). Gnathos
a broad transverse bar with anteriorly directed short
and narrow lateral arms. Valva 210—230 um, with very
long and slender apical process curved inwardly; inner
(ventral) lobe of valva extended into a distal process
with a few caudally directed spine-like outgrowths.
Transtilla absent (i.e., without transverse bar);
sublateral processes from valval bases short, slightly
curved outwards. Vinculum large; ventral plate trian-
gular, gradually narrowed anteriorly, without lateral
lobes. Aedeagus ca.354 um long, slightly broadened
towards base, with two spine-like cornuti, without
apical processes.

GENITALIA §. Not studied (see Remarks).

BIOLOGY. Adults collected in April.

DIAGNOSIS. Easily distinguished from other Enteucha
and other Neotropical nepticulids by the partially re-
duced tegumen and by the combination of large
triangular vinculum and very long and inwardly bent
distal process of the valva. This species is also well
characterized by other features that are not unique,
such as the whitish apex of the forewing, the huge
rounded uncus, and transverse bar-like gnathos.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL.

The holotype specimen is slightly mouldy; the other
known specimen (a female) is not included in the type
series.

DY;

MATERIAL EXAMINED.

Holotype 6, Belize: Cayo District, S of Teakettle

Village, Pook’s Hill Nature Reserve, 28—29.iv.1998

(Puplesis & Hill), genitalia slide no. 29117(BMNH).
Excluded from paratype series: 1 2, Belize: Chiquibul

Forest Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis &

Hill) (BMNB).

REMARKS. This species is named in honour of Vicki
and Ray Snaddon, owners of Pook’s Hill Nature Re-
serve, in recognition of their support for the
conservation of Belizean rainforest and its scientific
investigation.

E. snaddoni is morphologically interesting in that it
exhibits a small number of very marked modifications.
The partially reduced tegumen is functionally replaced
by the large oval uncus. The latter is surprisingly
unspecialized, however, although this is not to suggest
that is represents the nepticulid ground-plan. The trans-
verse form of the gnathos is very different from that of
most other nepticulids, and links this species with
gilvafascia; it is parallelled in a few other nepticulids
in Trifurcula and other genera.

The single female specimen available was collected
in a different area from the male holotype; no features
other than external similarity suggest that the two are
opposite sexes of the same species although we think
this is probable. As the association is doubtful, the
female is not included in the type series and its genita-
lia have been not documented.

MANONEURA Davis

7. Manoneura basidactyla (Davis, 1978),
comb. n.

(Figs 12, 61, 83-85, 207)

Oligoneura basidactyla Davis, 1978: 218-219.

MALE (Fig. 12). Forewing length: 1.7—2.1 mm. Wing-
span: 4.9 mm. Head: palpi yellowish cream to cream;
frontal tuft orange; collar a large tuft of lamellar yel-
lowish cream scales with gold lustre; eye-caps
yellowish cream; antenna brownish grey, ca. 38-39
segments. Thorax, tegulae and forewing fuscous brown
with bronze and very strong blue and purple lustre.
Forewing with distinct oblique yellow-gold subtermi-
nal fascia. Cilia fuscous, lighter distally. Underside of
forewing brown. Hindwing lanceolate, very slender,
dark brown with purple and blue lustre strongest on
upperside of hindwing; cilia brownish. No androconia
on forewing or hindwing. Legs grey except tarsi which
are cream. Abdomen blackish on upperside, blackish
grey on underside; genital segments blackish grey, not
contrasting with main colour of abdomen.

FEMALE. Similar to male. Forewing length: 2.3 mm.

R. PUPLESIS AND G.S. ROBINSON

Wingspan: 5.2 mm. Antenna ca. 32-33 segments. Legs
cream, shaded laterally with blackish. Abdomen black-
ish on upperside but cream on underside. Otherwise as
in male.

GENITALIA 6 (Figs 83-85). Capsule ca. 290-305 um
long. Tegumen with short, slightly bilobed, pseuduncus-
like caudal extension. Uncus with strong pointed
central process directed anteriorly and long narrow
lateral arms directed posteriorly. Gnathos with small
but complex and well-sclerotized v-shaped centre and
broad, membranous lateral arms. Process of uncus (in
natural position) almost touching posterior indenta-
tion at centre of gnathos, the structure appearing unified
(Fig. 84). Valva ca. 205-215 um, relatively narrow in
caudal half and gradually broadened towards base;
apical process long and slender. Transtilla absent, i.e.,
no transverse bar; however bases of valvae with re-
markably very long and straight sublateral processes.
Vinculum very long and broad, truncate at the anterior
end; no anterior emargination or lateral lobes. Aedeagus
236-246 um long, with two pairs of pointed lateral
carinae; no cornuti on vesica. Juxta with flask-shaped
outline, fused with aedeagus.

GENITALIA 2 (Fig. 207). Total length ca. 720 um.
Anal papillae undeveloped. S8 and T8 broadly rounded.
Apophyses posteriores short and complex. Apophyses
anteriores very slender and twice length of apophyses
posteriores. Vestibulum sclerotized. Caudal part of
corpus bursae very narrow caudally, swollen anteriorly;
no signa visible. Accessory sac undeveloped, repre-
sented by a small but clearly visible ring-like
sclerotization; ductus spermathecae long and narrow,
slightly sinuous.

BIOLOGY. Hostplant: Coccoloba uvifera (L.)
(Polygonaceae) (according to specimen label data
recorded by Davis, 1978: 218-219). Adults collected
in early January, April-May and July. The suggestion
by Davis (1978) that this species may be univoltine is
not supported by recently-collected material.

DIAGNOSIS. MM. basidactyla may be easily distin-
guished from the only other species in the genus, M.
trinaria (below) by the relatively straight valva and
anteriorly truncate vinculum; the two species are ex-
ternally similar in pattern and lustre, but basidactyla is
slightly paler. This species differs from the remaining
neotropical nepticulids by the strong purple lustre of
the forewing and the large anteriorly truncate vincu-
lum together with the distinctively-shaped gnathos.

DISTRIBUTION. Known from the southwestern coast
of Florida, Dominica (new record) and Belize (new
record). The species is likely to have a wide distri-
bution at least in the Caribbean region.

CONDITION OF TYPE MATERIAL. According Davis
(1978) two specimens from the four comprising the

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

type series are glued to paper points and one of these
glued specimens is lacking its abdomen. Holotype in
FSCA, paratypes in FSCA and USNM.

MATERIAL EXAMINED.

3d, 2°, Belize: Chiquibul Forest Reserve, Las Cuevas,
3-16.iv.1998 (Puplesis & Hill), genitalia slide nos.
291206, 29121 2 (BMNH); 26, 19, same data, wing
venation slide no. AD03142 (VPU). 16, Dominica:
Pont Casse, 16.v.1965 (Davis), genitalia slide no.
Diskus 002 (USNM); 16, 19, Cabrit Swam, 10-
13.v.1965 (Davis) (USNM); 2 exx. (without
abdomens), Springfield Est., 20—26.vii.1963 (Flint)
(USNM).

REMARKS. Thetwo species of this genus represent an
exotic and distinctive element of the Neotropical fauna.

8. Manoneura trinaria sp. n.
(Figs 13, 86, 87)

MALE (Fig. 13). Forewing length: 2.5 mm. Wing-
span: about 5.6 mm. Head: palpi cream; frontal tuft
orange; collar comprised of a single uniform tuft of
fuscous shiny lamellar scales; eye-caps cream; an-
tenna blackish grey, shiny, ca. 43 segments. Thorax,
tegulae and forewing fuscous with very strong purple
and blue lustre, which is especially intensive on
forewing; at certain angles forewing may have purple
and gold lustre instead of blue. Subterminal fascia
slightly curved inward, gold, shiny. Cilia fuscous-
brown, but tending to cream at tips. Underside of
forewing fuscous. Hindwing lanceolate, fuscous, at
certain angles with purple and bright grey lustre, espe-
cially along dorsal margin; cilia fuscous-brown. No
androconia visible on forewing or hindwing. Legs
fuscous, a little paler on tarsi. Abdomen fuscous on
upperside and underside, a pair of very short anal tufts
and genital segments also fuscous.

FEMALE. Unknown.

GENITALIA 6 (Figs 86, 87). Capsule 460-473 um
long. Tegumen extended into short rounded
pseuduncus-like sclerite with small lateral lobes. Un-
cus with relatively small strongly pointed central
process directed anteriorly and very long narrow lat-
eral arms directed posteriorly. Gnathos with small,
complex, well-sclerotized v-shaped centre and rather
membranous lateral arms. Process of uncus (in natural
position) almost touching posterior indentation at cen-
tre of gnathos, the structure appearing unified. Valva
about 170 um long, relatively narrow and distinctly
curved inward, without separate apical process and
with shorter, pointed dorsal lobe. Transtilla absent, no
transverse bar developed, but bases of valvae with long
and broad sublateral processes. Vinculum very long
and broad, narrowed in anterior half and very broadly
rounded anteriorly. Aedeagus 370-382 um long, with

23

two pairs of spine-like lateral carinae; no cornuti on
vesica. Juxta elongate, subrectangular, fused with
aedeagus.

BIOLOGY. Adults collected in January.

DIAGNOSIS. Externally this is one of the most strik-
ing nepticulid species by dint of its extremely strong
purple and blue forewing lustre; it differs from M.
basidactyla in the curved valva and long aedeagus and
vinculum. It differs from all other nepticulids of the
Neotropical region in the striking purple and blue
forewing lustre, the curved, distally divided valvae,
and long aedeagus with long straight apical carinae
with slightly globose bases.

DISTRIBUTION. Venezuela (cloud forest).

CONDITION OF TYPE MATERIAL. The single specimen
available is well-preserved, but the hindwings are ob-
scured beneath the unspread forewings, and it is
difficult to study them.

MATERIAL EXAMINED.

Holotype 6, Venezuela: Aragua [Rancho Grande],
1100 m, 17—20.1.1978 (Heppner), genitalia slide no.
Diskus003 (USNM).

REMARKS. This is one of the most striking repre-
sentatives of the Neotropical fauna with remarkable
forewing coloration, and with distinctively modified
gnathos and uncus (a feature shared only with
basidactyla). Phylogenetically both basidactyla and
trinaria are markedly apomorphic.

STIGMELLA Schrank

The salicis group
S. fuscotibiella-group Newton & Wilkinson, 1982.

The species described below are attributable on the
basis of forewing pattern and male genital structure to
the large salicis-group, first established for species of
the European fauna and known in the Nearctic as the
‘fuscotibiella-group’ (Newton & Wilkinson, 1982).
The term ‘salicis-group’ is used here for the Neotropical
species below to avoid further confusion. However, the
female genitalia of all the Neotropical species lack the
characteristic band (signum) of tiny, dentate chitin
plates encircling the bursa that is typical of the Holarctic
members of the group; just olyritis has a similar, but
most likely not homologous, structure on the bursa.
Thus the numerous Holarctic species appear to share a
synapomorphic character that is absent in the
Neotropical ones; in the latter the where plate-like
pectinations, if present, are scattered. The probability
of scattered or absent pectinations being the
apomorphic character-state seems intuitively less likely.

24

So the Holarctic representatives may form a mono-
phyletic entity, derived from a Neotropical-type
ancestor, within the salicis-group. Further speculation
on the geographical origin of the salicis-group, appar-
ently primitive and speciose in the Neotropics, is
possible but perhaps should await more thorough
exploration and understanding of the southern hemi-
sphere representatives.

9. Stigmella andina (Meyrick, 1915)
(Figs. 14, 88, 89, 209)

Nepticula andina Meyrick, 1915: 255, 256.
Stigmella andina (Meyrick); Davis, 1984: 18.

MALE (Fig. 14, left side). Forewing length: 2.4—
2.7 mm. Wingspan: 5.2—5.9 mm. Head: palpi cream;
frontal tuft cream or ochreous cream to dull ochre;
collar yellowish cream, indistinct; eye-caps pale cream
to yellowish cream; antenna brownish, ca. 34 seg-
ments. Thorax and tegulae pale cream with
yellow-ochre tints. Forewing uniformly glossy yel-
lowish cream with a brassy tint; no dark scales at apex
as in females; some whitish lustre at apex. Cilia whit-
ish cream to ochreous cream. Underside of forewing
uniformly brownish ochre or brownish. Hindwing lan-
ceolate, cream on both sides, cilia also cream. No
androconia on forewing or hindwing. Legs pale ochre-
ous cream or cream. Abdomen greyish brown on
upperside and underside, genital segments cream, dis-
tinctly contrasting in colour with rest of abdomen.

FEMALE (Fig. 14, right side). Head colour very simi-
lar to male; antenna a little shorter (ca. 30 segments).
Apical one-quarter to one-fifth of forewing entirely
covered with purple brown scales, some brownish
scales overlapping on cilia. Abdomen ochreous cream,
sometimes pale ochre. Slender ochre-brown oviposi-
tor distinctly visible at end of abdomen. Otherwise as
in male.

GENITALIA 6 (Figs. 88, 89). Capsule 370 um long.
Uncus relatively long, well-sclerotized, with four dis-
tinct caudal papillae. Tegumen short, strip-like, with
caudal concavity, strongly sclerotized and rounded
laterally. Gnathos with well-developed central plate
and two long posterior processes; lateral arms of
gnathos narrow in lateral view. Valva 208 um, broad,
with short, bent apical process, which may not be very
evident in lateral view. Transtilla curved laterally, with
broad transverse bar, but without sublateral processes.
Vinculum relatively broad, anterior margin with deep
semicircular emargination and pointed triangular lat-
eral lobes. Aedeagus 242 um long, without apical
processes. Vesica with about 10 very large, caudally-
pointed cornuti (the number of cornuti is usually
significant in this group).

R. PUPLESIS AND G.S. ROBINSON

GENITALIA @ (Fig. 209). Total length 1180 um. S8
narrow, but rounded. Anal papillae not developed. T8
with a few small setae not segregated into lateral
groups. Apophyses posteriores long (ca. 370 um) and
very broad, broadly curved anteriorly. Apophyses
anteriores slender, as long as apophyses posteriores,
curved slightly inwards anteriorly. Vestibulum rela-
tively narrow, without pectinations or sclerites. Caudal
part of corpus bursae very broad and strongly folded;
remaining larger part of corpus bursae broadly oval,
without distinct pectinations or signa. Accessory sac
small, but broad; ductus spermathecae slender and
slightly sinuous.

BIOLOGY. Adults collected in July.

DIAGNOSIS. Belongs to the S. salicis-group. The
broad, rather square valva with a tiny, curved apical
process together with 10 very large vesical cornuti
distinguishes this species from all other Neotropical or
world representatives of the salicis-group. There are
slight superficial resemblances to the Nearctic S.
fuscotibiella (Clemens) but it differs in having large
vesical cornuti and a pale forewing. See Remarks.

DISTRIBUTION. Peru.

CONDITION OF TYPE MATERIAL. Most specimens are
in poor condition, badly pinned and many without
head scales or even lacking a head; the forewing pat-
tern is barely visible, and sometimes very worn. The
lectotype and two other specimens are in compara-
tively satisfactory condition.

MATERIAL EXAMINED.
Lectotype d, Peru: [Oroya], 12200 ft [3720 ml,
vul.1914 (Parish), genitalia slide no. 20612 (BMNH),
here designated.

Paralectotypes, 13d, 6°, data as lectotype (genitalia
slide no. 28841 9(BMNH).

REMARKS. According to the original description of
this species (Meyrick, 1915), sexual dimorphism is
characteristic (Fig. 14). However, there is no guarantee
that the female paralectotypes really belong to the
same species as the male lectotype. Differences in
male and female forewing coloration may represent
outstanding sexual dimorphism or simply indicate two
different species. Work with a sample of unidentified
rubbed material has shown that seven externally very
similar specimens collected in adjacent areas of the
Peruvian Andes may represent seven different species
of Stigmella. So the association of males and females
of this species should be treated with caution.

10. Stigmella cuprata (Meyrick, 1915)
(Figs 15, 90, 91)
Nepticula cuprata Meyrick, 1915: 255.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE
Stigmella cuprata (Meyrick); Davis, 1984: 18.

MALE (Fig. 15). Forewing length: 1.9 mm. Wing-
span: 4.3 mm. Head: palpi greyish cream; frontal tuft
yellowish; collar yellowish, consisting of piliform
scales; eye-caps relatively small, whitish with metallic
lustre; antenna brownish grey, ca. 32 segments. Tho-
rax, tegulae and basal half of forewing before fascia
greyish bronze. Medial fascia of forewing silvery,
glossy, broad, proximal margin slightly oblique, distal
margin broadly rounded. Forewing beyond fascia purp-
lish brown with a few apical scales just before cilia
which may have silver lustre at certain angles. Cilia
grey. Underside of forewing brown. Hindwing and
cilia brownish grey. No androconia on hindwing or
forewing.

FEMALE. Not studied (see Remarks).

GENITALIA 6 (Figs90,91). Total length 280-286 pm.
Uncus trapezoidal with a pair of small rectangular
caudal lobes and a pair of tiny lateral papillae. Tegumen
small with bulged lateral corners. Gnathos with
well-developed central plate, two-long close-set cau-
dal processes and narrow sclerotized lateral arms. Valva
about 185 pm, triangular, gradually narrowed into a
pointed, inward-curved apical process. Transtilla with
strong ribbon-like transverse bar but without sublateral
processes. Vinculum with moderately large lateral
lobes, which gradually narrow anteriorly; anterior
emargination semicircular. Juxta absent. Aedeagus
290-305 um. Vesica with about 11 huge cornuti and a
few minute and inconspicuous thorn-like cornuti par-
ticularly on left side subapically.

BIOLOGY. Adults collected in July.

DIAGNOSIS. Belongs to the S. salicis-group. Exter-
nally resembles S. johannis (Zeller), but the male
genitalia differ strongly in the closely set caudal proc-
esses of the gnathos, the caudally narrowed bilobed
uncus, and large lateral lobes of the vinculum. From
other species of the group, including the Nearctic S.
slingerlandella (Kearfott), cuprata differs in the broad
silvery medial fascia and by the group of very large
cornuti in the aedeagus.

DISTRIBUTION. Peru.

CONDITION OF TYPE MATERIAL. The left antenna of
the lectotype is broken, the right forewing rubbed. The
other specimen in the type series (probably @) has
badly rubbed forewings and is badly pinned.

MATERIAL EXAMINED.
Lectotype 6, Peru: Matucana, 7780 ft [2370 ml],
vii.1914 (Parish), genitalia slide no. 28848 (BMNH),
here designated.

Paralectotype ??, data as holotype.

REMARKS. Both specimens of the original syntype

25

series are present in the BMNH collection. We have
redefined this taxon from the male lectotype. The
other specimen (probably a female) is extremely
rubbed, badly pinned, and with few or no traces of
pattern. It is impossible to tell whether it is likely to be
conspecific with the lectotype and we have therefore
not dissected it. No external features remain that are
worth describing.

11. Stigmella johannis (Zeller, 1877)
(Figs 16, 92, 93)

Nepticula johannis Zeller, 1877: 456-457.
Stigmella johannis (Zeller); Davis, 1984: 18.

MALE (Fig. 16). Forewing length: about 2.3 mm.
Wingspan: about 5.4 mm. Head: palpi whitish cream;
frontal tuft very pale orange; collar cream, slightly
darkened with brownish tint, consisting of lamellar
scales; eye-caps pale cream; antenna grey-brown, ca.
36 segments. Thorax, tegulae and forewing smooth
brown with a gold tint. Postmedian fascia of forewing
silvery, lustrous, not clearly defined. Cilia brownish.
Some scales of forewing close to cilia with silvery
lustre at certain angles. Underside of forewing intense
brown with purple lustre along entire length. Hindwing
and cilia grey or brownish. No androconia on hindwing
or forewing. Legs grey-brown. Abdomen grey-brown
above and below; genital segments cream.

FEMALE. Unknown.

GENITALIA 6 (Figs 92, 93). Capsule 315-320 um.
Uncus subtrapezoidal, slightly tapered caudally, with
four relatively large caudal papillae. Tegumen small,
laterally slightly bulged. Gnathos with relatively small
central plate and large, well-separated, almost parallel
posterior processes; lateral arms moderately devel-
oped. Valva 215-225 um, subtriangular, with two small,
pointed, inwardly-curved apical processes. Transtilla
with transverse bar exceeding anterior margin of vin-
culum; sublateral processes of transtilla absent, corners
broadly rounded. Vinculum with small pointed trian-
gular lateral lobes; anterior emargination shallow,
almost square. Juxta absent. Aedeagus about 340 um,
with about nine well-sclerotized large cornuti and a
smaller less sclerotized one.

BIOLOGY. The single adult known was collected in
late May.

DIAGNOSIS. Belongs to the S. salicis-group. Differs
from other representatives of the group (including the
Neotropical S. cuprata (Meyrick) and Nearctic S.
slingerlandella (Kearfott)) by the combination of lus-
trous silvery postmedian fascia, many almost
equal-sized large cornuti in the aedeagus, well-sepa-
rated caudal processes of the gnathos, and broad uncus
with four caudal papillae.

26
DISTRIBUTION. Colombia.

CONDITION OF TYPE MATERIAL. ‘The holotype is in
satisfactory condition with clear wing pattern, but the
wings are not spread.

MATERIAL EXAMINED.
Holotype 3, Colombia: Bogota, 27.v.1871 (Johann),
genitalia slide no. 28843 (BMNH).

12. Stigmella rudis sp. n.
(Figs 17, 94-96, 208)

MALE (Fig. 17). Forewing length: 2.9—3.8 mm. Wing-
span: 6.6—8.3 mm. Head: palpi grey to greyish cream
or cream; frontal tuft pale orange or yellow; collar
comprising two tufts of whitish lamellar scales; eye-
caps whitish; antenna brown to dark brown, ca. 36
segments. Thorax and tegulae brown or dark brown.
Forewing densely irrorated with brown, dark brown
and some grey-cream scales; with two irregular and
variable greyish cream postmedian spots of which the
costal may be bigger than the tornal. Cilia grey to
greyish cream distally. Underside of forewing brown.
Hindwing lanceolate, relatively broad, grey to brown;
cilia concolorous. No androconia on forewing or
hindwing. Legs with tinted brown or dark grey. Abdo-
men fuscous brown on upperside and underside; genital
segments cream (distinctly contrasting in colour with
remaining abdomen) to occasionally grey.

FEMALE. Colour similar to male; antenna shorter (ca.
29 segments).

GENITALIA 6 (Figs 94-96). Capsule 346-356 um
long. Uncus relatively long, well-sclerotized, with four
distinct caudal papillae and oval caudal emargination.
Tegumen very short, simple, band-shaped. Gnathos
with narrow central plate and two very long almost
parallel posterior processes, with a pair of broad ante-
rior extensions and narrow and relatively long lateral
arms. Valva 235-245 um long, very broad, almost
square, with two distinct pointed apical processes.
Transtilla with slender transverse bar forming short
triangular sublateral lobes. Vinculum relatively short,
with broad, square anterior emargination and with
anteriorly rounded short triangular lateral lobes.
Aedeagus 268-287 um long, without apical processes.
Vesica with about 7 spine-like cornuti and with a
highly characteristic large cluster of variably-sized
spine-like cornuti, most of them fused at the base.
Juxta not apparent in the studied preparations.

GENITALIA 2 (Fig. 208). Total length ca. 1117 pm.
S8 and T8 truncate caudally. Anal papillae not devel-
oped. Apophyses posteriores short and very slender in
anterior one-third to one-fifth. Apophyses anteriores
very slender and very long (ca. 235 um). Vestibulum
relatively broad, membranous, without pectinations or

R. PUPLESIS AND G.S. ROBINSON

sclerites. Caudal section of corpus bursae broad and
strongly folded; remaining larger part of corpus bursae
broadly oval, entirely covered with distinct scattered
irregular pectinations. Accessory sac moderately de-
veloped, more or less oval and folded; ductus
spermathecae slender, weakly sclerotized and without
convolutions.

BIOLOGY. Adults common from October to Febru-
ary.

DIAGNOSIS. Belongs to the S. salicis-group. Recog-
nisable by the unique cluster of spine-like cornuti
arising from a fused base. The species is also charac-
terized by the somewhat featureless forewing pattern
which separates it from other Neotropical representa-
tives of the group.

DISTRIBUTION. Chile and Argentina.

CONDITION OF TYPE MATERIAL. Most specimens (in-
cluding the holotype) are in satisfactory or good
condition, a few are rubbed.

MATERIAL EXAMINED.

Holotype 3d, Argentina: Rio Negro: San Carlos de
Bariloche, Colonia Suiza, 800 m, 7—30.x1i.1981
(Nielsen & Karsholt), genitalia slide no. Diskus175
(ZMUC).

Paratypes, Argentina: 9 d, data as holotype, genita-
lia slide no. Diskus180 (ZMUC); 14, Lago Nahuel
Huapi, Puerto Blest, 770 m, 22—31.xii.1981 (Nielsen
& Karsholt) (ZMUC); 36, Neuquen, San Martin de
Los Andes, 640 m, 17.x—26.x1.1981 (Nielsen &
Karsholt) (ZMUC); 3, 4°, Lago Lacar, Pucara, 650—
750 m, 25.xi.1978 & 28—29.xi.1981 (Mission Cientifica
Danesa and Nielsen & Karsholt), genitalia slide no.
Diskus176 2(ZMUC); 1d, Lago Lacar, 5 km E of Hua-
Hum, 640 m, 25.xi.1981 (Nielsen & Karsholt)
(ZMUC); 3d, 1 2, Chubut, Esquel, Lago Menendez, El
Sagrario Puerto, 550-600 m, 21.11.1979 and 2-4.1.1982
(Mission Cientifica Danesa and Nielsen & Karsholt),
genitalia slide no. Diskus205 2 (ZMUC). Chile: 7d,
2 2 Osorno, Parque National Puyehue, Aguas Calientes,
450 m, 10.xii.1981 (Nielsen & Karsholt), genitalia
slide nos. DiSkus177 9, Diskus179¢ (ZMUC); 14,
Anticura, 350 m, 19.xi.1981 (Nielsen & Karsholt)
(ZMUC).

REMARKS. Externally this species varies in forewing
length and pattern: some specimens may be smaller
than average or darker. Genital structures, especially
the highly characteristic cluster of cornuti in the
aedeagus represent lees variable features and are most
useful for recognition of the taxon.

13. Stigmella marmorea sp. n.
(Figs 18, 97, 98, 210)

MALE (Fig. 18). Forewing length: 4.04.6 mm. Wing-

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

span: 9.0-10.1 mm. Head: palpi brownish; frontal tuft
comprised of whitish cream and dark brown piliform
scales; collar forming two clearly separated tufts of
whitish or cream lamellar scales; eye-caps whitish or
cream; antenna dark grey-brown, ca. 43-44 segments.
Thorax and tegulae dark grey-brown except for grey-
ish margins. Forewing brown with distinctive marbled
pattern (hence the species’ name) comprising wide
cream or whitish antemedian and postmedian fasciae;
some whitish scales between fasciae, and many just
before the cilia that are so numerous that they form a
narrow terminal fascia. Cilia brown-grey to greyish at
tornus. Underside of forewing grey-brown. Hindwing
lanceolate, relatively broad, grey; cilia grey. No
androconia on forewing or hindwing. Legs intensely
shaded with grey-black or grey-brown, tarsi black on
one side. Abdomen dark grey on upperside, cream on
underside; genital segments cream, not contrasting
with underside of abdomen but clearly distinguishable
because of large valval lobes.

FEMALE. Externally similar to male.

GENITALIA ¢ (Figs 97, 98). Capsule 478-489 um
long. Uncus relatively long, well-sclerotized, with
four slender caudal papillae and deep, oval caudal
emargination. Tegumen very short, with rounded lat-
eral angles, bulged in the middle. Gnathos with
moderately narrow central plate and two very long
parallel posterior processes; a pair of anterior exten-
sions of the gnathos are very tiny, not always distinct;
lateral arms of gnathos narrow, relatively short. Valva
305-312 pm long, almost equally broad throughout
entire length or gradually narrowed towards apex, with
two close-set pointed apical processes. Transtilla with
slender transverse bar forming short triangular lobe-
like sublateral processes. Ventral plate of vinculum
half or less valval length, with broad anterior
emargination and almost pointed triangular lateral
lobes. Aedeagus 351-365 um long, without apical
processes but with serrated oblique apical margin.
Vesica with about 9-10 large or very large horn-like
cornuti and with a group of a few smaller spine-like
cornuti at apex; one cornutus is huge, one-half length
of aedeagus. Juxta distinct, triangular, narrowing
anteriorly.

GENITALIA ?(Fig.210). Total length ca. 894-907 um.
S8 and T8 broadly rounded caudally. Anal papillae not
developed. Apophyses posteriores just a little shorter
than apophyses anteriores which are long (ca. 233—
241 um.) and very slender. Vestibulum narrow,
membranous, without pectinations or sclerites. Cau-
dal region of corpus bursae broad and folded; remainder
relatively small, comprising only half bursa length,
with sparse and scattered pectinations. Accessory sac
well-developed, folded; ductus spermathecae slender,
weakly sclerotized and without convolutions.

27
BIOLOGY. Adults collected in March.

DIAGNOSIS. Belongs to the S. salicis-group. Differs
from other species, including Neotropical ones, in
having two white or cream fasciae on the forewing, and
an apically serrated apical margin on the aedeagus
which contains one huge cornutus among numerous

others.
DISTRIBUTION. Peru (altitudes around 4100 m).

CONDITION OF TYPE MATERIAL.
served.

Relatively well-pre-

MATERIAL EXAMINED.
Holotype d, Peru: Dept. Ancash, 23 km SE of Huaraz,
Cerro Cahuish [Quabrada Pucavado], 4100 m, 15—
18.1i1.1987 (Karsholt), genitalia slide no. Diskus 182d
(ZMUC).

Paratypes, 1d, 1 9, data as holotype, genitalia slide
no. Diskus181 2 (ZMUC).

14. Stigmella peruanica sp. n.
(Figs. 19, 104, 105)

MALE (Fig. 19). Forewing length: ca. 2.1 mm. Wing-
span: 4.6 mm. Head: palpi grey-brown; frontal tuft
pale orange; collar comprising two clearly separated
tufts of fuscous grey lamellar scales with gold lustre;
eye-caps cream or yellowish cream; antenna fuscous,
ca. 32—33 segments. Thorax and tegulae fuscous with
strong bronze-gold lustre. Forewing with three shiny
fasciae: (1) moderately large, dark yellow-gold basal
fascia, (2) wide whitish gold median fascia, (3) very
narrow and short whitish gold terminal fascia; areas
between fasciae with smoothly dark brown scales with
gold and purple lustre. Cilia fuscous brown. Underside
of forewing fuscous. Hindwing lanceolate, relatively
broad, grey-brown; cilia concolorous. No androconia
on forewing or hindwing. Legs dark grey with metallic
lustre. Abdomen grey-brown on upperside, grey to
ochreous cream on underside; short anal tufts grey-
brown; genital segments ochreous cream, not
particularly contrasting in colour with underside of
abdomen.

FEMALE. Unknown.

GENITALIA 6 (Figs 104, 105). Capsule 312-325 um
long. Uncus relatively long, well-sclerotized, with four
distinct caudal papillae and deep oval caudal
emargination. Tegumen very short, simple, band-like.
Gnathos with large medially narrowed central plate
and two slender, elongate posterior parallel processes;
anterior extensions triangular; lateral arms tapering
caudally, very short. Valva 204—210 um long, slightly
narrowed apically, with two close-set pointed apical
processes. Transtilla with strong transverse bar, with-
out sublateral processes. Ventral plate of vinculum
relatively short, half or less valval length; anterior

28

emargination broad but shallow, lateral lobes small,
anteriorly subtruncate. Aedeagus broad (98—112 tum)
and relatively short (256-269 um), without apical proc-
esses. Vesica with about 10 large or very large horn-like
cornuti, four of them very long and positioned cen-
trally and towards base, six (or more) about half that
length and in a row caudally from one-half. Juxta
triangular, with shallow longitudinal division at slen-
der caudal end, rather membranous, broad anterior
margin indistinct.

BIOLOGY. Adults collected in March.

DIAGNOSIS. Easily recognisable among all other
species of the group by the three gold lustrous mark-
ings on the forewing and the “4 + row of 6’ configuration
of cornuti in the aedeagus.

DISTRIBUTION. Peru (altitudes around 3000 m).

CONDITION OF TYPE MATERIAL. Holotype in rela-
tively good condition, however the left forewing is
slightly damaged.

MATERIAL EXAMINED.
Holotype 6, Peru: Dept. Puno, 5 km E of Limbani,
3000 m,

28.11.1987 (Karsholt), genitalia slide no. Diskus 189
(ZMUC).

15. Stigmella epicosma (Meyrick, 1915)
(Figs 20, 21, 99-103, 211, 212)

Nepticula epicosma Meyrick, 1915: 255.
Stigmella epicosma (Meyrick); Davis, 1984: 18.

MALE (Fig. 20). Forewing length: 1.8—2.0 mm. Wing-
span: 4.1—4.4 mm. Head: palpi brownish cream; frontal
tuft yellowish or orange-yellow; collar large, creamy
white, comprised of lamellar scales (see Remarks);
eye-caps relatively small, whitish, distinctly glossy;
antenna grey-brown to blackish, ca. 29-32 segments.
Thorax and tegulae brown with gold lustre. Forewing
variable, with either two or three glossy silver fasciae
with some indistinct gold lustre: (1) postmedian (some-
times median), broad and slightly oblique; (2) terminal,
narrow, following the wing margin just before cilia,
tending to be interrupted in the middle; (3) basal, can
be present or absent — if present (as in 2 shown in
Fig. 21) then narrower than median fascia, if absent
then forewing base is a little lighter (Fig. 20) (see
Remarks). Ground colour between fasciae brown to
dark brown with gold lustre. Cilia brown distally.
Underside of forewing brown. Hindwing and cilia
brownish to brown or grey. No androconia on hindwing
or forewing. Legs predominantly darkened with brown.
Abdomen grey-fuscous above and below; short anal
tufts fuscous; genitalia segments (predominantly val-
vae) visible from underside, ochreous-cream,
contrasting with main colour of abdomen.

R. PUPLESIS AND G.S. ROBINSON

FEMALE (Fig. 21). Very similar to male.

GENITALIA 6 (Figs 99-103). Capsule 290-317 pm.
Tegumen small, caudally slightly bulged. Uncus with
two well-sclerotized papillate lobes, distinctly sepa-
rated by deep medial emargination. Gnathos with very
long, slender, almost parallel caudal processes, narrow
lateral arms and short, wide central plate, slightly
emarginate anteriorly. Valva 170—195 um, broad, with
inwardly curved and pointed apical process. Transtilla
with transverse bar (which may be narrow or broad)
and very broad, lobate sublateral processes. Valvae
with additional slender basal connection. Ventral plate
of vinculum relatively short but broad, with weakly
developed and anteriorly broadly rounded lateral lobes
or with well-developed triangular lobes; anterior
emargination from very shallow and indistinct to semi-
circular. Aedeagus 195—220 um, with only two huge
cornuti (Fig. 102). Occasionally the left cornutus may
be half the length of that illustrated (see Remarks).

GENITALIA @ (Figs 211, 212). Total length about
600-645 um. Anal papillae weakly developed, two
wide but very short setose lobes. Apophyses anteriores
almost straight, curved inward anteriorly. Apophyses
posteriores very slender and long, their anterior ends
not exceeding apophyses anteriores. Corpus bursae
relatively small and narrow, no signa or pectinations
visible. Accessory sac folded, not distinctly demar-
cated from vestibulum; ductus spermathecae
membranous, except for semicircular sclerotization at
a distance from accessory sac.

BIOLOGY. Adults collected in February, April and
August.

DIAGNOSIS. Belongs to the S. salicis-group. Distin-
guished from all other Stigmella by the glossy silver
fasciae of the forewing and by the presence only two,
but very large, cornuti in the aedeagus. The presence of
two huge cornuti in the aedeagus is also a feature of
two other Neotropical species (schoorli and hamata)
but these do not have glossy silver median (or postme-
dian) and apical fasciae on the forewing.

DISTRIBUTION.
4000 m).

Peru (widely distributed, 150-

CONDITION OF TYPE MATERIAL. ‘The paralectotype
(d) has the wings unspread, but is in satisfactory
condition; the lectotype has a broken left antenna; it
and the female paralectotype have the wings crudely
spread but in the latter the forewings and head scaling
are abraded.

MATERIAL EXAMINED.

Lectotype d, Peru: Lima, 150 m [‘500 ft’], viii.1914
(Parish), genitalia slides no. 288496 (BMNH), here
designated.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

Paralectotypes, 1d, 1 9, data as lectotype, genitalia
slide no. 28850 2(BMNH).

Non-type material: Peru: 7d, 2 2, Dept. Arequipa,
8 km E Arequipa, Rio Andamayo Valley, 2920 m,
7.iv.1987 (Karsholt), genitalia slide nos Diskus 186 &,
Diskus 187d (ZMUC); 164, Dept. Lima, 10 km N of
Oyon, Quabrada Quichas Pueblo Quichas, 4000 m,
24—26.ii.1987 (Karsholt), genitalia slide no. Diskus
188 (ZMUC).

REMARKS. The type series of three specimens (la-
belled as collected at 500 ft) do not have the basal
fascia developed; the basal area has a stronger gold
lustre, looks paler, and a few weakly silver-lustred
glossy scales can be seen on one specimen. All speci-
mens in a recently-collected series from 2920 m in
Peru have a distinct glossy silver basal fascia. One
(possibly aberrant) male specimen collected at 4000 m
has the basal area of the forewing glossy gold, a
fuscous-gold shiny collar and greyish white palpi. The
genitalia of this specimen (slide no. DiSkus 188,
ZMUC) are typical for the species, but the left cornutus
in the aedeagus is half the length of that in a normal
specimen (Fig. 102 — normal). These differences in
coloration and minor details of genital structure prob-
ably represent geographical variation in mountainous,
dissected terrain.

16. Stigmella schoorli sp. n.
(Figs 24, 106, 107, 213, 214)

MALE (Fig. 24). Forewing length: 2.5—2.7 mm. Wing-
span: 5.8-6.0 mm. Head: palpi cream; frontal tuft
comprised of dark brown and ochre cream piliform
scales; collar forming pair of broad, clearly separated
tufts of cream or golden cream lamellar scales; eye-
caps whitish to golden cream; antenna dark grey to
blackish grey, with gold lustre, ca. 35-36 segments.
Thorax, tegulae and forewing uniformly dark grey
with strong bronze and gold lustre; a few scales just
before forewing cilia may appear glossy white at a
certain angles. Cilia grey to dark grey. Underside of
forewing dark brown with light purple lustre. Hindwing
and cilia grey. No androconia on forewing or hindwing.
Legs cream or grey-cream, shaded laterally with black-
ish grey. Abdomen fuscous.

FEMALE. Forewing length: 2.7—2.8 mm. Wingspan:
5.8-6.5 mm. Antenna ca. 26-27 segments. Similar to
male.

GENITALIA 6 (Figs 106, 107). Capsule 340-364 um
long. Uncus long, well-sclerotized, with two relatively
long parallel quadrate lobes, each with a few indistinct
papillae; caudal emargination of uncus u-shaped.
Tegumen very short, band-like. Gnathos with moder-
ately narrow but well-sclerotized central plate and two

29

very long parallel posterior processes; pair of anterior
extensions very small, not always distinct; lateral arms
narrow and relatively short. Valva 221—232 um long,
broad, with sinuous inner side and large pointed apical
process; inner lobe of valva forming broad triangular
subapical extension. Transtilla with strong transverse
bar and huge lobe-like sublateral processes. Ventral
plate of vinculum less than half length of valva, with
very shallow and broad anterior emargination; lateral
lobes weakly developed, broadly rounded anteriorly.
Aedeagus 232-256 um long, without apical processes,
relatively broad (99-106 um). Vesica with very distinc-
tive pair of very large parallel cornuti; one about 159
um long, the other 128 um long. Juxta triangular,
membranous, indistinct, slightly bifid at apex.

GENITALIA @ (Figs 213, 214). Total length about
750 um. S8 and T8 almost truncate caudally. Anal
papillae not developed. Apophyses anteriores a little
shorter than apophyses posteriores, which are slender
and long (ca. 166 um.). Vestibulum narrow, membra-
nous, folded, without pectinations or sclerites. Caudal
half of corpus bursae broad and folded; anterior half
comparatively small, swollen anteriorly, without vis-
ible pectinations or signa. Accessory sac
well-developed, large, elongate and folded; ductus
spermathecae slender, without convolutions but with
an irregularly shaped sclerotization 200 um from the
accessory sac.

BIOLOGY. Adults collected in February.

DIAGNOSIS. Belongs to the S. salicis-group. The most
distinctive feature of the species — the presence of only
two very large cornuti — is shared only with the
Neotropical epicosma and hamata. From the first
schoorli may be easily distinguished by the uniformly
coloured forewing, and from the second by the short
and broad aedeagus, long sublateral transtilla proc-
esses, distinctly bilobed uncus, long apical process of
the valva and weak anterior emargination of the vincu-
lum. Because of the external similarities between
schoorli and hamata these species may not be reliably
separated except by dissection of the genitalia. How-
ever, schoorli differs slightly differs from S. hamata
externally: specimens are distinctly smaller, the frontal
tuft is a mix of dark grey and ochreous cream scales (in
hamata the scales are uniformly coloured), the
forewings are notas gold and glossy, and the hindwings
are darker.

Although belonging to the same species group and
occurring in the same geographical area, having very
similar external features, and sharing a similar distinc-
tive pattern of cornuti, schoorli and hamata are not
phylogenetically close and represent distinct and rec-
ognizable species.

DISTRIBUTION. Peru (3870 m).

30

CONDITION OF TYPE MATERIAL. Mostly good.

MATERIAL EXAMINED.
Holotype d, Peru: Dept. Ancash, 35 km SE of Huaraz,
Cerro Cahuish [Quabrada Pucavado], 3870 m,
18.iii. 1987 (Karsholt), genitalia slide no. Diskus 2003
(ZMUC).

Paratypes: 3d, 5, data as holotype, genitalia slide
nos. Diskus 201—203 6, Diskus 204-206 2 (ZMUC).

REMARKS. This species is named in honour of Dr J.W.
Schoorl (Amsterdam).

17. Stigmella hamata sp. n.
(Figs 22, 108-112)

MALE (Fig. 22). Forewing length: 3.1—3.3 mm. Wing-
span: 6.8—7.0 mm. Head: palpi greyish brown; frontal
tuft brownish ochre, collar and eye-caps greyish cream,
glossy; antenna grey with gold lustre, ca. 38 segments.
Thorax, tegulae and forewing uniformly grey with
strong gold lustre; a few scales just before forewing
cilia may appear glossy and whitish at certain angles;
cilia brownish to brownish cream at tornus. Underside
of forewing grey. Hindwing and cilia brownish. No
androconia on forewing or hindwing. Legs cream with
shaded with brown and grey. Colour of abdomen un-
known.

FEMALE. Unknown.

GENITALIA 6 (Figs 108-112). Capsule ca. 365 um
long. Uncus long, well-sclerotized, tapered caudally,
with four distinct caudal papillae. Tegumen short,
bulged in the middle, lateral angles rounded. Gnathos
with moderately narrow, but well-sclerotized central
plate and two very long and slender parallel posterior
processes; without anterior extensions; lateral arms
narrow, moderately long. Valva 260-272 um long,
almost equally broad throughout length, with pair of
broad, inturned apical processes. Transtilla with short
transverse bar, angles rounded and slightly extended.
Ventral plate of vinculum short, with v-shaped anterior
emargination; lateral lobes more or less triangular,
rounded anteriorly. Aedeagus 312—324 um long, with-
out apical processes, narrow (47—58 pm). Vesica with
three horn-like cornuti; one of them very large, about
95 um, another at apex half this length, the third
shorter, often obscured by the first. Juxta membranous
and indistinct, probably triangular.

BIOLOGY. Adults collected in March.

DIAGNOSIS. Belongs to the S. salicis-group. Charac-
terized by the uniformly grey forewings with strong
gold lustre, and aedeagus with only two large cornuti
(of three). Externally resembles schoorli which also
occurs in the Peruvian Andes. However, hamata is a
notably larger and with a uniformly coloured ochreous

R. PUPLESIS AND G.S. ROBINSON

frontal tuft, stronger gold lustre of forewing, paler
hindwing and very distinctive male genitalia: the un-
cus has four caudal papillae whereas in schoorli it is
bilobed; the aedeagus is narrower and the valva has
distinctive broad hook-shaped apical processes.

DISTRIBUTION. Peru (3240 m).

CONDITION OF TYPE MATERIAL. Specimens of the
type series are well-preserved.

MATERIAL EXAMINED.

Holotype 3, Peru: Dept. Cuzco, 40 km NW Sicuani,

5 km E of Laguna Pomacanchi, 3240 m, 24.11.1987

(Karsholt), genitalia slide no. Diskus 199d (ZMUC).
Paratype 6, Peru: Dept. Puno, 15 km E of Ayaviri,

1d, 26-27.iii.1987 (Karsholt), genitalia slide no.

Diskus198d (ZMUC).

18. Stigmella imperatoria sp. n.
(Figs 23, 113-116)

MALE (Fig. 23). Forewing length: 4.1-4.3 mm. Wing-
span: 9.0-9.6 mm. Head: palpi cream to greyish or
grey, distally and on underside usually fuscous; frontal
tuft very large, comprised of cream or pale ochreous
and numerous dark brown piliform scales, the latter
mostly central; collar of mainly cream or greyish cream
lamellar scales, additionally with some overlapping
piliform scales; eye-caps small, almost covered by
frontal tuft, glossy greyish cream with a few piliform
scales anteriorly; antenna blackish brown with gold
and purple lustre, ca. 54 segments. Thorax, tegulae and
forewing uniformly brown with strong copper-gold
lustre, no purple. Cilia brown. Underside of forewing
dark grey-brown. Hindwing and cilia brown. No
androconia on forewing or hindwing. Legs cream with
fuscous brown or fuscous grey lateral shading. Abdo-
men grey-brown on upperside, yellowish cream on
underside; genital segments paler but not contrasting
significantly with main colour of abdominal sternites.

FEMALE. Unknown.

GENITALIA 6 (Figs 113-116). Capsule 669-685 um
long. Uncus unusually long, very strongly sclerotized
laterally, slightly broadened at apex and with two large
lateral papillae and two weakly-developed central pa-
pillae; caudal emargination of uncus (usually present
in related species) absent. Tegumen very short, simple,
band-shaped. Gnathos with narrow central plate and
two very long, almost parallel posterior processes with
apices turned outwards; anterior extensions of gnathos
undeveloped, but anterior margin with shallow central
emargination; lateral arms narrow and moderately long.
Valva ca. 434 um long, relatively narrow and of uni-
form breadth, with large pointed inturned apical
process; inner lobe of valva forming broad triangular
subapical extension. Transtilla with short and rather

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

slender transverse bar characterized by pointed and
anteriorly-directed angles, but without sublateral proc-
esses. Ventral plate of vinculum large, with v-shaped
anterior emargination and large, anteriorly rounded
lateral lobes. Aedeagus extremely long (756-767 um )
and slender (ca. 92-100 um), without apical processes.
Vesica with a apical group of about 10 large spine-like
cornuti and with a long basal band of mainly very tiny
needle-like or spine-like cornuti mixed with just a few
larger ones. Juxta a simple, weakly sclerotized plate
broadening towards caudal end.

BIOLOGY. Adults collected in February.

DIAGNOSIS. Belongs to the S. salicis-group. Easily
distinguishable from all currently known representa-
tives of this group by the distinctive long and narrow
uncus and aedeagus, and also by the copper-gold lustre
of forewing. The configuration of the cornuti in the
aedeagus is diagnostic and distinctive.

DISTRIBUTION. Peru (4100 m).

CONDITION OF TYPE MATERIAL. Specimens, includ-
ing the holotype, are well-preserved; the hindwings of
one paratype have been broken off and placed in a
small plastic capsule below the specimen.

MATERIAL EXAMINED.
Holotype d, Peru: Dept. Ancash, 35 km SE of Huaraz,
Cerro Cahuish [Quabrada Pucavado], 4100 m, 15—
18.11.1987 (Karsholt), genitalia slide no. DiSkus195d
(ZMUC).

Paratypes, 2d, data as holotype, genitalia slide no.
Diskus196d (ZMUC).

19. Stigmella olyritis (Meyrick, 1915)
(Figs 28, 131, 132)

Nepticula olyritis Meyrick, 1915: 256
Stigmella olyritis (Meyrick); Davis, 1984: 18.

MALE (Fig. 28). Forewing length: 1.7—1.9 mm. Wing-
span: 3.84.2 mm. Head: palpi brownish white; frontal
tuft white; collar rather indistinct, white, comprised of
piliform scales; eye-caps small, whitish, lustrous; an-
tenna brown. Thorax and tegulae brown. Forewing
irrorated with grey-brown, brownish cream and cream
scales; basal half of forewing more smoothly scaled
than apical half, where pale scales are especially abun-
dant on costal and tornal margins. Fascia absent. Cilia
brownish to yellowish cream, brown at tornus. Under-
side of forewing brown. Hindwing and cilia brown. No
androconia on hindwing or forewing. Legs grey-brown.

Abdomen brown.
FEMALE. Similar to male.

GENITALIA 6 (Figs 131, 132). Capsule 250-256 pm.

31

Uncus almost semicircular, with numerous caudal
papillae. Tegumen short, simple, anteriorly
well-sclerotized and sinuous. Gnathos with moder-
ately developed central plate and very large
convergent posterior processes; lateral arms small.
Valva 158-160 um long, tapering, with two slender
and close-set apical spine-like processes. Transtilla
with slender but well-sclerotized transverse bar;
sublateral corners of transtilla extended into weakly
demarcated triangular processes. Vinculum long with
small triangular and anteriorly pointed lateral lobes;
anterior emargination shallow, broadly semicircular.
Aedeagus long, about 280 um. Vesica with numerous
dense spine-like cornuti mostly collected into one
band, additionally with very tiny spine-like or trian-
gular cornuti distributed around the main band. Juxta
present, a simple quadrate plate.

GENITALIA 2? (Fig. 215). Total length 958-960 um.
Narrowly constricted S8 and T8 forming a distinctly
slender and rather long ovipositor. Apophyses
posteriores very slender and long (320-325 um),
slightly divergent at anterior end. Apophyses anteriores
slightly longer than apophyses posteriores, very slen-
der in anterior half. Vestibulum narrow and folded.
Caudal part of corpus bursae broad and strongly folded,
remaining anterior region very large, spheroidal or
near-spherical, with inverted U-shaped signum with
numerous pectinations along it (see Remarks). Acces-
sory sac almost spherical, small but well-developed;
ductus spermathecae helical with about 8.5 turns.

BIOLOGY. Adults collected in August.

DIAGNOSIS. Belongs to the salicis species-group. The
caudally almost rounded uncus together with the very
long, close-set gnathos processes, the short sublateral
processes of the transtilla, long vinculum and numer-
ous subequal cornuti unequivocally distinguish this
species from all other members of the group. The
external features of olyritis are unfortunately much
less helpful for immediate diagnosis.

DISTRIBUTION. Peru.

CONDITION OF TYPE MATERIAL. This species was
described by Meyrick from six specimens (3 d, 3 9); all
are in unsatisfactory condition, badly pinned and some-
what abraded.

MATERIAL EXAMINED.
Lectotype d, Peru: Lima, 150 m [‘500 ft’], viii.1914
(Parish), genitalia slide no. 288516 (BMNH), here
designated.

Paralectotypes: 2d, 3 9, data as holotype, genitalia
slide no. 28852 2? (BMNH).

REMARKS. ‘The signum in the female genitalia does
not look convincingly homologous with that of
Holarctic representatives of the group; it is similar in

32

shape but not in structure, being comprised of dense
pectinations.

The eurydesma group

This group is newly established in the Nepticulidae for
species with a unique autapomorphic configuration of
gnathos and uncus: the gnathos has a pair of short,
lateral, horn-like, caudally directed processes close up
against the uncus, and a reduced central plate; the
processes are joined by a weakly sclerotized, almost
invisible anterior connection. The uncus is a small
quadrate plate with or without a caudal emargination,
partially occluded by the gnathos and the lateral regions
difficult to view; the base appears to be partly fused
with the bases of the gnathos processes. Species of this
group are also distinguished by the valva being broad,
and tapered towards the apex, with a well-developed
pointed apical process; the aedeagus has numerous
fine spine-like cornuti; a juxta, as far as is known, is
always present and generally plate-shaped. The
forewing is fuscous with a distinct median or post-
median fascia.

The group slightly resembles a few Nearctic
Stigmella (such as tiliella Braun) in a few aspects of
morphology but differs in the partially reduced gnathos
and more elaborated uncus. A close phylogenetic rela-
tionship between these species and _ the
eurydesma-group 1s possible, but further investigation
is needed to support this. The group is possibly end-
emic to the Neotropical region.

20. Stigmella eurydesma (Meyrick, 1915)
(Figs 25, 128-130)

Nepticula eurydesma Meyrick, 1915: 255
Stigmella eurydesma (Meyrick); Davis, 1984: 18.

MALE (Fig. 25). Forewing length: about 1.6 mm.
Wingspan: 3.6-3.7 mm. Head: palpi cream; frontal
tuft ochreous orange; collar distinctly broad, com-
posed of pale cream lamellar scales; eye-caps pale
cream; antenna brown to fuscous brown, ca. 20 seg-
ments. Thorax and tegulae dark brown. Forewing
fuscous brown with some purplish lustre which is most
distinctive apically; fascia median — postmedian, very
broad, pale cream, tending to broaden dorsally. Cilia
fuscous or brown. Underside of forewing dark brown.
Hindwing lanceolate, brown, cilia brown to grey-
brown. No androconia on forewing or hindwing. Legs
mainly ochreous cream shaded with grey-brown.
Abdomen fuscous brown, genital segments dark grey
brown, very weakly contrasting in colour with remain-
der of abdomen.

FEMALE. Forewing length: about 1.7 mm. Wingspan:
about 3.8 mm. Number of antennal segments unknown.

R. PUPLESIS AND G.S. ROBINSON

Forewing fascia usually slightly broader than in males,
distinctly broadening dorsally. Hindwing brown to
pale ochre-brown. Abdomen fuscous brown on
upperside, grey-brown to ochre-brown on underside.
Otherwise as male.

GENITALIA 6 (Figs 128-130). Capsule 264 um long.
Uncus small, with two well-sclerotized, quadrate lobes.
Tegumen short, well-sclerotized posteriorly. Gnathos
comprising two lateral horn-like processes, so strongly
sclerotized as to appear black in a preparation, and
with very weakly developed, almost membranous ant-
erior transverse bar (probably representing a remnant
of central plate); posteriorly directed processes of
gnathos very close to uncus lobes. Valva 176 um,
pointed and strongly sclerotized apically, but very
weakly sclerotized and specialized in basal one-third.
Transtilla with well-developed sublateral processes
and narrow, but well-sclerotized transverse bar; trans-
verse bar connects sublateral processes not at their
anterior extremities, as is usual in Stigmella, but half
way along the length of the processes. Vinculum short,
with shallow anterior emargination and small lateral
lobes. Aedeagus ca.250 pm long, without apical proc-
esses. Vesica with numerous more or less equally long
spine-like cornuti concentrated towards apex and nu-
merous very tiny spine-like cornuti concentrated
towards base. Juxta present, represented by a simple
plate narrowed at middle between weakly sclerotized
bases of valvae.

GENITALIA °. Not studied because of very poor con-
dition of available material (see Remarks).

BIOLOGY. Adults collected in April.

DIAGNOSIS. Externally this species could be con-
fused with other Stigmella species with a white
postmedian fascia. However, the shape of gnathos, in
which the central plate is greatly reduced separates the
species from all currently known representatives of
Stigmella except for the related albilamina and
fuscilamina from which it differs in the by weakly
sclerotized bulged inner side of the valva and by the
absence of very large or plate-shaped cornuti.

DISTRIBUTION. Guyana.

CONDITION OF TYPE MATERIAL. Although the wings
of the lectotype are not well spread and the specimen is
slightly rubbed, structures and forewing pattern are
more or less well-preserved. The condition of the
paralectotypes is very poor. One male is in a gelatine
capsule (head structures and wing pattern
well-preserved); two female specimens lack the head
and are very poorly pinned; the remaining female has
a head, but the abdomen is lacking.

MATERIAL EXAMINED.
Lectotype 6, Guyana: Georgetown, iv.1913 (Parish),

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

genitalia slide no. 28842 (BMNH), here designated.
Paralectotypes: 1d, 3 9, data as lectotype (BMNH).

REMARKS. No material additional to Meyrick’s type
series is known. The female genitalia of specimens
from the type series have been left undissected as they
are extremely fragile and very poorly pinned. We
believe that removal of an abdomen cannot be accom-
plished without very high risk of damage. This is not
commensurate with the potential gain in information.
Although Acalyptris females do show considerable
interspecific variation in genital morphology, we do
not expect great differences between those of
eurydesma, albilamina and fuscilamina. Studies of
females of the latter two (which we cannot separate or
unequivocally associate with males) show that the
abdominal tip forms a short but strongly sclerotized
ovipositor, both apophyses are long and slender, the
accessory sac is oval and neatly folded, the folded part
of the bursa copulatrix is long, slender and densely
folded, and the corpus bursae proper is small with
indistinct spinose pectinations.

21. Stigmella albilamina sp. n.
(Figs 26, 63, 117-122)

MALE (Fig. 26). Forewing length: 1.9—2.2 mm. Wing-
span: 4.3-5.0 mm. Head: palpi white; face from wholly
white to brown to blackish closer to tuft; frontal tuft
pale to dark ochre, large; collar large, creamy white,
comprised of lamellar scales; eye-caps creamy white,
large; antenna black, ca. 23 segments. Thorax and
tegulae brown-black. Forewing brown-black to black,
occasionally dark brown, with weak, occasionally dis-
tinct, purplish lustre and postmedian, occasionally
distinctly postmedian, whitish fascia; proximal margin
of fascia usually transverse, sometimes slightly con-
cave; distal margin oblique and therefore fascia usually
broader on dorsum; fascia occasionally narrow. Cilia
grey-brown, paler distally. Underside of forewing fus-
cous with weak purplish lustre. Hindwing and cilia
from brown to grey-brown. Basal one-half to two-
fifths of hindwing with grey androconial scales which
are broad but short, overlapping nearly one-fifth length
of cilia on dorsal margin, however in most specimens
they are indistinct or indiscernible. No androconia on
forewing. Legs pale cream on underside, usually black-
ish or grey on upperside or laterally. Abdomen
brown-black on upperside but with distinctly contrast-
ing cream genital segments at tip; underside cream, at
least in posterior half, genital segments (valvae) vis-
ible as two broad rounded or triangular plates wholly
covered with pale cream scales (in fuscilamina they are

predominantly fuscous, in eurydesma — grey-brown).
FEMALE. Not studied (see Remarks).

GENITALIA 6 (Figs 117-122). Capsule 268-270 um.

33

Uncus a small quadrate plate, laterally strongly
sclerotized with slightly extended corners, and with
medial emargination. Gnathos with pair of short, blunt,
lateral caudally-directed processes very close to un-
cus; processes well-sclerotized (but not black as in
fuscilamina); central plate of gnathos reduced, lateral
processes jointed by weakly sclerotized, almost invis-
ible anterior connection. Valva 195 um long, relatively
broad, with sclerotized inner margin and slightly
inturned apical process. Transtilla with broad sublateral
process and transverse bar. Juxta strongly sclerotized,
simple, quadrate, always distinct between valvae. Ven-
tral plate of vinculum very small, narrowed in the
centre, with very short and minute pointed lateral
lobes; anterior emargination very shallow, broad and
slightly rounded. Aedeagus 305-330 um, without api-
cal carinae; vesica with numerous small and only
weakly sclerotized spine-like cornuti and one huge
horn-shaped cornutus (occasionally two); oval sclerite
(present in fuscilamina) absent.

BIOLOGY. Adults collected in April; very common.

DIAGNOSIS. The shape of gnathos, with the central
plate strongly reduced, separates this species from all
Stigmella except the related eurydesma and
fuscilamina. From these two albilamina differs in the
degree of sclerotization of the gnathos which looks
dark, but not black as in eurydesma, in the short lateral
lobes of the vinculum, and the presence of one huge
cornutus in addition to numerous weakly developed
spine-like cornuti (in eurydesma, and particularly in
fuscilamina they are bigger and more strongly
sclerotized). From eurydesma this species differs also
in the well sclerotized inner margin of the valva, the
shape of the transtilla and juxta (not narrowed in the
middle); from fuscilamina it differs in that the vincu-
lum lobes are much shorter, the inner surface of the
valva is less sclerotized, there is no plate-like cornutus
(and there is no huge horn-like cornutus in fuscilamina),
the juxta is distinctive, the uncus has a medial
emargination and produced corners, there are some-
times visible androconia on the hindwing, and the
genital segments are pale cream, the valvae appearing
as a pair of whitish lamina (fuscous in fuscilamina).

DISTRIBUTION. Belize.

MATERIAL EXAMINED.
Holotype 3, Belize: Chiquibul Forest Reserve, Las
Cuevas, 3-16.iv.1998 (Puplesis & Hill), genitalia slide
no. 29113 (BMNH).

Paratypes, 58d, data as holotype, genitalia slide nos
29112 (BMNH), AD0306 (VPU) (BMNH (49); VPU
(9)).

REMARKS. The male genitalia of several additional
specimens have been studied as temporary prepara-
tions in glycerin — these are not listed above. There is

34

what we presume to be a long mixed series of female
albilamina and fuscilamina in BMNH, but at present
we are unable to satisfactorily distinguish two species
among them or to correctly associate females with
males. Until this species is reared the identity of the
female remains problematic and we have not described
the genitalia. [See Remarks for eurydesma.]

S. albilamina and fuscilamina are closely related
sister-species occurring in the same habitat and having
coincident flight times. Nevertheless they, or at least
the males, are morphologically distinct. Light trapping
in Belize in April 1998 showed albilamina to be some-
what more common than fuscilamina.

22. Stigmella fuscilamina sp. n.
(Figs 27, 123-127)

MALE (Fig. 27). Forewing length: 2.0—-2.2 mm. Wing-
span: 4.8-5.0 mm. Head: palpi white, occasionally
shaded laterally with black; face brown to black, but
usually white close to tuft; frontal tuft pale to dark
ochre, large; collar large, creamy white, comprised of
lamellar scales; eye-caps creamy white, large; antenna
black, sometimes pale at apex or underside, ca. 23-26
segments. Thorax and tegulae brown-black. Forewing
brown-black to black with weak purplish lustre and
with median to postmedian whitish fascia; proximal
margin of fascia transverse, the distal margin oblique,
thus fascia usually broader at dorsum; fascia occasion-
ally narrow. Cilia grey-brown, paler distally, sometimes
almost cream. Underside of forewing fuscous with
weak purplish lustre. Hindwing and cilia brown to
grey-brown. No androconia on hindwing or forewing.
Legs whitish cream on underside, usually black or
grey-brown on upperside and laterally. Abdomen
brown-black on upperside and underside; genitalia
segments visible on underside as two broad rounded or
broadly triangular plates, valvae covered with
brown-black scales, cream only at apex (in contrast to
albilamina, where the entire valva is covered with
whitish scales).

FEMALE. Not studied (see Remarks).

GENITALIA 6 (Figs 123-127). Capsule 268-270 um.
Uncus a small quadrate plate with rounded corners and
without caudal emargination. Tegumen small,
band-shaped. Gnathos with two short, horn-like,
caudally-directed lateral processes close-set to uncus;
processes very strongly sclerotized, appearing black in
preparations, overlapping lateral margins of uncus;
central plate of gnathos reduced, lateral horn-like proc-
esses joined by weakly sclerotized, almost invisible
anterior connection. Valva 178-195 um long, rela-
tively broad, with distinctly sclerotized inner margin,
apical process curved inward. Transtilla strongly and
conspicuously sclerotized laterally, with moderately

R. PUPLESIS AND G.S. ROBINSON

long and slender sublateral process and long trans-
verse bar. Juxta barely visible, represented by a
membranous and caudally bilobed plate. Ventral plate
of vinculum relatively small, with long, slender,
spine-like lateral lobes; anterior emargination quad-
rate. Aedeagus 305-317 um; vesica with numerous
moderately large and strongly sclerotized spine-like
cornuti and large oval plate-like sclerite.

BIOLOGY. Adults collected in April.

DIAGNOSIS. The shape of the gnathos, with the cen-
tral plate strongly reduced, separates this species from
all currently known representatives of Stigmella ex-
cept related eurydesma and albilamina. From these
two it differs in the long lateral lobes of the vinculum,
and the presence of a big plate-like cornutus in addi-
tion to numerous well-developed spinose cornuti (in
eurydesma all cornuti are spinose, in albilamina they
are weakly developed). It differs also in the sclerotized
inner side of the valva, and almost invisible membra-
nous juxta. It differs from eurydesma in the shape of
the transtilla and the almost straight caudal margin to
the juxta; from albilamina it differs in the vinculum
lobes being three times as long, the black gnathos,
significantly more strongly sclerotized inner side of
the valva, the absence of a huge horn-like cornutus and
the remaining cornuti being bigger and more strongly
sclerotized. The uncus in fuscilamina, in contrast to
albilamina, is not emarginated caudally and has
rounded, rather that sharp corners; there are no
androconia on the hindwing whereas these are some-
times visible in albilamina. The genital segments are
fuscous-scaled — if a specimen is examined without
dissection there are two clearly fuscous laminae vis-
ible — these are the valvae. In albilamina the valvae are
cream.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL.
well-preserved.

Most specimens are

MATERIAL EXAMINED.
Holotype, 6, Belize: Chiquibul Forest Reserve, Las
Cuevas, 3—16.i1v.1998 (Puplesis & Hill), genitalia slide
no. 29110 (BMNH).

Paratypes, 18d, data as holotype, genitalia slide nos
28111 (BMNH), AD0305 (VPU) (BMNH (12); VPU (6)).

REMARKS. The male genitalia of several additional
specimens have been studied as temporary prepara-
tions in glycerin — these are not listed above. Although
we have what we take to be a mixed series of females
of this species and albilamina we are unable to differ-
entiate them or associate them with males so we have
not described the genitalia. [See also Remarks for
eurydesma.|

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

Species unattributed to a group

23. Stigmella gossypii (Forbes & Leonard,
1930)

(Figs. 29, 133, 134)

Nepticula gossypii Forbes & Leonard, 1930: 149.

Stigmella gossypii (Forbes & Leonard); Newton &
Wilkinson, 1982: 404, 405, figs 32, 33; Davis,
1984: 18.

MALE (Fig. 29). [Adapted from Newton & Wilkinson,
1982.] Forewing length: 3.0 mm. Wingspan: about
6.5 mm. Head: palpi whitish; frontal tuft ochre; collar
white; eye-caps creamy white, lustrous; antenna dark
brown, number of segments unknown. Thorax and
tegulae grey with metallic lustre. Forewing dark brown
with purple and silver lustre, with shiny gold
antemedian fascia and narrower, shiny silver postme-
dian fascia. Cilia greyish with metallic grey sheen,
contrasting with row of pale wing scales. Hindwing
lanceolate, grey, cilia shiny metallic grey. No
androconia are described for this species (see Rem-
arks). Legs brown, shiny metallic grey on lower surface.
Abdomen dark brown on upperside, shiny metallic
grey on underside.

FEMALE. Similar to male, but tending to be a little
larger (forewing about 3.0-3.2 mm) (Newton &
Wilkinson, 1982).

GENITALIA 6 (Figs 133, 134). [Adapted from New-
ton & Wilkinson, 1982.] Capsule ca. 228 um long.
Uncus quadrate, corners slightly extended and pointed.
Tegumen a small transverse bar. Gnathos broad
U-shaped with a narrow transverse bar, very tiny ante-
rior processes and long slender posterior processes.
Valva 146 um long, gradually narrowed towards apex
and bifid apically. Transtilla with well-developed
sublateral processes and narrow transverse bar, which
is not fused in the middle. Vinculum relatively small
with distinct pointed lateral lobes; anterior
emargination semicircular. Juxta a trapezoidal mem-
branous plate, broadening slightly towards caudal end.
Aedeagus 230 um long, without apical processes.
Vesica with very numerous and very small spinose
cornuti.

GENITALIA @ [Adapted from Newton & Wilkinson,
1982 and their fig. 33.] Total length 338 um. S8 mod-
erately broad. Anal papillae weakly developed, very
short and broadly rounded. T8 rounded posteriorly.
Apophyses posteriores long and slender (ca. 64 um).
Apophyses anteriores slender and almost as long as
posterior ones. Vestibulum membranous, without
scobination. Caudal part of corpus bursae relatively
narrow, with few very indistinct comb-like pectinations;

35

remaining large and ovoid, scobination rather indis-
tinct and represented by numerous comb-like
pectinations. Accessory sac large, almost two-fifths
length of corpus bursae, slightly folded transversely;
ductus spermathecae slender and probably not convo-
luted.

BIOLOGY. According Forbes & Leonard (1930) lar-
vae (sometimes in great number) mine leaves of
Gossypium barbadense, occasionally G. hirsutum. Egg
on underside of leaf, in a fork of small leaf-veins. Mine
slender, sinuous, mainly on underside of leaf, more
than 25 mm in straightened length. Frass granular,
deposited in arcuate waves in first half of mine, in a
continuous line in latter half. Mines on a single leaf
may cross and anastomose frequently. Mines are almost
invisible if the upperside of fresh leaves is examined.
When leaves become old and dry the mines become
whitish in marked contrast to the dark leaf surface.
(Forbes & Leonard, 1930; Newton & Wilkinson, 1982).
Larva green with pale brownish head, 2.5 mm. Cocoon
white, oval, flattened, strongly narrowed to posterior
end, ca. 1.8 x 1.0 mm, on underside of leaf. Forbes &
Leonard, (1930) suggested that there was only one
generation per year but the voltinism of this species is
not proven. Adults fly in late March and April.

DIAGNOSIS. Slightly resembles a few Nearctic
Stigmella species (prunifoliella (Clemens), ceanothi
(Braun), rhoifoliella (Braun), heteromelis Newton &
Wilkinson, diffasciae (Braun)) which also possess a
broad U-shaped gnathos. However, gossypii differs
from all similar species in the combination of distally
divided valva, slender gnathos process, long pointed
lobes of vinculum and two broad shiny forewing fas-
ciae, the proximal fascia gold and the distal one silver.

DISTRIBUTION. Puerto Rico; USA (Florida).

MATERIAL EXAMINED.

None. The holotype (2) and 6 paratypes are deposited
in USNM and were studied by Newton & Wilkinson
(1982); they were not re-examined by us.

24. Stigmella kimae sp. n.
(Figs 30, 135-138)

MALE (Fig. 30). Forewing length: about 1.7 mm.
Wingspan: about 4.2 mm. Head: palpi cream; frontal
tuft and eye-caps almost unicolorous, pale cream;
collar tending to be a little darker, but still whitish;
antenna dark brown, about 26 segments. Thorax,
tegulae and forewing blackish brown with gold gloss
and some greenish and purple lustre. Single median-
postmedian fascia of forewing snow-white. Cilia
brownish, well-defined at apex by row of brown la-
mellar wing scales. Underside of forewing fuscous.
Hindwing and cilia grey. No androconia visible on
forewing or hindwing. Legs ochreous grey, shaded

36

fuscous. Abdomen relatively small, intense fuscous on
upperside and underside, genital segments (valval
lobes) small, brownish cream, contrasting in colour
with remainder of abdomen.

FEMALE. Unknown.

GENITALIA 6 (Figs 135-138). Capsule 233-252 um
long. Uncus with quadrate lateral lobes. Tegumen band-
like. Gnathos broad U-shaped, with long sinuous
posterior processes connected via slender but long
transverse bar (central plate); anterior processes of
gnathos very short, slightly bent inwards. Valva ca.
146 um long, oval, with very short almost blunt apical
process, which is only weakly separated from bulged
inner lobe of valva. Transtilla with short and slender
sublateral processes and long slender transverse bar.
Vinculum very short, without anterior emargination or
lateral lobes. Aedeagus 287—296 um long, rather broad
(ca.110 um at broadest point), without apical proc-
esses. Vesica with numerous tiny spine-like cornuti
and an apical group of about five large horn-like cornuti.

BIOLOGY. Adults collected in April.

DIAGNOSIS. Related to the Nearctic filiella (Braun),
but differing in the snow-white forewing fascia (shiny
silver in ftiliella), paler hindwing, whitish frontal tuft
(ochreous in filiella); the male genitalia primarily dif-
fer in the presence of sublateral processes on the
transtilla and the single cluster of long horn-like cornuti
in aedeagus. There are no other species closely related
to kimae. See Remarks.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. ‘The wing pattern in
the holotype is well-preserved and not abraded, but the
specimen is badly pinned and very fragile.

MATERIAL EXAMINED.

Holotype d, Belize: Chiquibul Forest Reserve, Las
Cuevas, 3—16.1v.1998 (Puplesis & Hill), genitalia slide
no. 29118 (BMNH).

REMARKS. Stigmella kimae and the Nearctic tiliella
(Braun) represent two close and distinctive species
among other Stigmella: both have rounded valvae, the
gnathos broadly U-shaped, the uncus with quadrate
well-separated lobes, very short vinculum, and the
aedeagus with one or two apical clusters of spine-like
cornuti. The relationships of the two are otherwise
obscure; they might be related to the Palaearctic S.
prunetorum species-group or to the widely distributed
luteella or ultima species-groups. The discovery of
further related species could prompt the recognition of
a separate species-group to contain these distinctive
taxa. Stigmella tiliella is known from Ohio and Ken-
tucky (USA) and is a bivoltine leaf-miner of Tilia
americana producing a long linear mine with a ten-
dency towards a spiral form; frass is deposited

R. PUPLESIS AND G.S. ROBINSON

irregularly across the entire breadth of the mine in the
first four-fifths but as a dense, central line in the
terminal portion (Newton & Wilkinson, 1982).

25. Stigmella plumosetaeella Newton &
Wilkinson, 1982

(Figs 31, 144, 145, 216)

Stigmella plumosetaeella Newton & Wilkinson, 1982:
455-456.

MALE (Fig.31). Forewing length: 1.6—-1.8 mm. Wing-
span: 3.4—3.9 mm. Head: palpi cream to greyish; frontal
tuft pale orange to pale ochreous; collar large, clearly
divided into two lateral tufts of lamellar scales, cream
or yellowish cream; eye-caps yellowish cream to shiny
white (see Newton & Wilkinson, 1982: 455); antenna
ochreous cream to brownish, ca. 27—28 short seg-
ments. Thorax and tegulae dark brown. Forewing base
and apical area beyond fascia brown to dark brown or
occasionally to orange-ochre or ferruginous; single
median fascia very broad, gradually broadening tow-
ards dorsal margin, yellowish or yellowish cream.
Cilia and some lamellar wing scales before cilia yel-
lowish or yellowish cream. Underside of forewing
grey to brown with some purple and some weak green
lustre. Hindwing and cilia brownish or grey, some-
times even smooth cream, depending upon illumination
and angle of view. No androconial patches visible on
forewing or hindwing. Legs cream with brown shading
laterally. Abdomen brownish or brownish cream on
upperside, cream or yellowish cream on underside;
distinct anal tuft cream, distinct; genital segments not
contrasting in colour with rest of abdomen.

FEMALE. Antennaca. 19-21 segments; otherwise as
in male (including the lustre of the forewing under-
side).

GENITALIA 6 (Figs 144, 145). Capsule 316-331 um
long. Uncus large, rounded caudally, with small caudal
emargination in centre or deeply bilobed (see Newton
& Wilkinson, 1982: 455, fig. 82). Tegumen small,
band-shaped. Gnathos with two long and slender pos-
terior processes connected via slender and long
transverse bar (central plate) and short anterior proc-
esses which may be rather broad and triangular. Valva
209-220 um long, reaching well beyond uncus, with
two pointed apical lobes and with long plumose scales
in apical half. Transtilla with distinct, lobate triangular
sublateral processes and transverse bar which is not
fused in the middle. Vinculum broad, almost
trapezoidal, without anterior emargination or lateral
lobes; if anterior margin concave then only very slightly
so. Aedeagus 220-268 pum long, gradually or abruptly
broadened in apical two-fifths, without apical proc-
esses. Vesica with numerous tiny spine-like cornuti at

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

apex of aedeagus. Juxta absent, but there are small
membranous lateral lobes on aedeagus.

GENITALIA @ (Fig. 216). Total length 562-645 um.
Abdominal tip broadly rounded. Anal papillae unde-
veloped. Apophyses posteriores long (ca. 116 um) and
slender. Apophyses anteriores half as long, slender in
anterior one-quarter to one-third. Vestibulum broad
and wrinkled. Caudal part of corpus bursae very slen-
der, slightly folded; remaining larger part of corpus
bursae relatively small, subspherical, entirely or almost
entirely lined with tiny pointed spines. Accessory sac
(according to Newton & Wilkinson, 1982) large and
covered with small denticles, but in Mexican speci-
mens an accessory sac is either absent or indiscernible,
and the ductus spermathecae is long, slender and sinu-
ous.

BIOLOGY. Adults collected in July and August.

DIAGNOSIS. Externally and in features of the male
genitalia this is a very distinctive species, character-
ized by the broad cream forewing fascia, relatively
slender, apically split valvae with very long plumose
setae, laterally rounded uncus, almost square ventral
plate of the vinculum and apically broadened aedeagus
with numerous very tiny cornuti. The presence of
plumose setae on the valva suggests a possible affinity
with the Neotropical S. barbata or with some Nearctic
representatives of the Stigmella saginella-group (see
Newton & Wilkinson, 1982). However, the specialized
shape of the valva, transtilla, uncus and apically broad-
ened aedeagus isolate plumosetaeella from these taxa.

DISTRIBUTION. USA (Arizona); Mexico.

CONDITION OF TYPE MATERIAL. The holotype male
and paratype female were deposited in ANS following
their description and illustration by Newton &
Wilkinson (1982); we did not re-examine them.

MATERIAL EXAMINED.

12d, 79, Mexico: 4 and 6 miles S of C. Victoria,
6.viii. 1963 (Duckworth & Davis), genitalia slide nos
Diskus004 3, Diskus005 d, DiSkus006 2? (USNM).

REMARKS. We follow Newton & Wilkinson (1982)
in regarding this species as occupying a very isolated
taxonomic position. However, further detailed revi-
sion of the Nearctic saginella-group may indicate
relationship between plumosetaeella, barbata (below)
and a few Nearctic species currently belonging to the
saginella-group.

26. Stigmella barbata sp. n.
(Figs 32, 139-143)

MALE (Fig. 32). Forewing length: 1.9—2.0 mm. Wing-
span: 4.4-4.5 mm. Head: palpi cream; frontal tuft
brown; collar and eye-caps whitish; antenna brown, ca.

37

22-23 segments. Thorax and tegulae brown. Forewing
brown with some indistinct golden lustre; a few sparse
brownish cream scales at apex of forewing and numer-
ous long thickened brown androconial scales on the
dorsal margin overlapping hindwing. Cilia and under-
side of forewing brown. Hindwing brown, with long
and relatively broad brown androconial scales in basal
two-thirds overlapping one-third length of scales of
cilia; cilia brown. Legs cream with brown shading.
Abdomen grey-brown on upperside, brownish cream
on underside, a distinctive pair of relatively long cream
anal tufts directed anteriorly; genital segments grey-
cream, not contrasting in colour with ventral surface of
abdomen.

FEMALE. Unknown.

GENITALIA 6 (Figs 139-143). Capsule ca. 244 um
long. Uncus distinctly bilobed, lobes triangular, turned
slightly outwards. Tegumen band-shaped, slightly
bulged caudally. Gnathos with two slender and rela-
tively long posterior processes connected via slender
transverse bar (central plate); anterior processes half or
one-third length of posterior processes; lateral arms
slender, distinct, almost same length as anterior proc-
esses. Valva about 195 um long, with broad,
inward-turned apical process and broad, bulged inner
lobe; inner side of apical process with long plumose
setae. Transtilla with short triangular sublateral proc-
esses and very slender transverse bar, which does not
fuse in the middle. Vinculum very short, ventral plate
trapezoidal, without anterior emargination or lateral
lobes. Aedeagus about 170 um long, without apical
processes, simple, slightly swollen at base. Vesica with
two strongly sclerotized apical spine-like cornuti which
appear fused at their bases, and a few tiny weakly
sclerotized cornuti; ventral side of aedeagus with nar-
row band-shaped sclerotization just beyond middle of
tube, apex of aedeagus finely and bluntly serrated.
Juxta broad at base, inconspicuous caudally.

BIOLOGY. Adults collected in April.

DIAGNOSIS. A remarkable species characterized by
elongate, plumose setae on the apical process of the
distinctly rounded valva; a brown frontal tuft, long
androconial scales on the forewing dorsum and
hindwing anal margin. The plumose valval scales and
similarities in the shape of the uncus, transtilla,
aedeagus and valva suggest this species may possibly
be attributed to the Neotropical saginella-group (origi-
nally designated by Wilkinson & Scoble, 1979).
However, the strongly bulged inner lobe of the valva,
and the slender and well-separated caudal processes of
the gnathos make the diagnosis of barbata uncompli-
cated. S. barbata differs from plumosetaeella (which
was not included in the saginella-group by Newton &
Wilkinson (1982)) in the brown forewing, the simple
aedeagus, bulged valva and well-separated uncus lobes.

38

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The single specimen
is relatively well-preserved but the frontal tuft is
abraded.

MATERIAL EXAMINED.

Holotype 6, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 29137 (BMNH).

REMARKS. In spite of similarities to the Nearctic
saginella-group, this species cannot be attributed to
this group without detailed revision of all the Nearctic
representatives.

27. Stigmella pruinosa sp. n.
(Figs 33, 34, 146, 147, 218)

MALE (Figs 33[ right side], 34). Forewing length:
1.5-1.7 mm. Wingspan: 3.3-3.7 mm. Head: palpi
cream; frontal tuft usually pale yellow to yellow-orange,
occasionally pale orange-ochre; collar whitish or pale
cream; eye-caps pale cream to (occasionally) yellow-
cream; antenna brown to dark brown (occasionally
pale brown) but brownish cream or whitish on basal
part of flagellum and underside, ca. 19-20 segments.
Thorax and tegulae brown, sometimes dark brown.
Forewing relatively narrow, brown to dark brown with
weak gold (not in all examples), occasionally bronze
lustre; sometimes with very weak purplish lustre.
Forewing with very numerous long and slender grey-
white androconial scales on upperside, densely
covering entire median area of wing and extending
across the width; size of androconial patch may vary
considerably from one-third to one-half length of
forewing, but base and apex of wing always lack
androconia. Cilia brown, usually paler at tips. Under-
side of forewing brownish, without androconial scales.
Hindwing brownish (or grey) with very distinct
androconial patch on basal one-third to two-fifths
(Fig. 34); oval androconial patch comprised mainly of
deep black scales, but with numerous elongate whitish
androconia scattered along middle of patch. Black and
whitish androconial scales overlap cilia for one-fifth of
their length. Cilia brown. Underside of hindwing
brown, without androconia. Legs cream with brown to
dark grey lateral shading. Abdomen brown to dark
brown on upperside, brownish cream on underside;
anal tufts indistinct, cream; scaling of genital segments
(mainly valvae) brownish cream.

FEMALE (Fig. 33, left side). Antenna ca. 18 seg-
ments. Forewing entirely brown, without greyish white
androconial scales; generally paler than in males, but
with some overlap — a few females are darker than the
palest males. Hindwing without androconial patch,
entirely brown. Otherwise similar to male.

R. PUPLESIS AND G.S. ROBINSON

GENITALIA ¢ (Figs 146, 147). Capsule 230 um long.
Uncus with two strongly sclerotized, quadrate lateral
lobes and U-shaped caudal emargination between the
lobes. Tegumen band-shaped, slightly extended
caudally. Gnathos with two posterior and two anterior
processes on well-developed relatively broad trans-
verse bar (central plate); anterior processes one-third
longer than posterior ones and twice as widely spaced;
lateral arms short. Valva ca. 122 um long, gradually
narrowed posteriorly, with small pointed, inturned api-
cal process. Transtilla with well developed transverse
bar, rounded laterally, without sublateral processes.
Vinculum relatively large, with broad emargination
anteriorly and with triangular lateral lobes. Aedeagus
very large, considerably longer than genital capsule
(ca. 307 um), without apical processes, slightly bent at
middle. Vesica with numerous tiny spine-like cornuti
and long row of considerably larger cornuti, the latter
weakly sclerotized and inconspicuous.

GENITALIA @ (Fig. 218). Total length ca.717 pm.
Abdominal tip narrow. Apophyses posteriores and apo-
physes anteriores almost same length (ca.
130-160 pm), but apophyses anteriores more slender.
Vestibulum narrow. Caudal part of corpus bursae broad,
slightly wrinkled; remaining larger part of corpus bur-
sae elongate, without pectination or signa. Accessory
sac rounded, weakly separated from bursa; shape of
ductus spermathecae unknown.

BIOLOGY. Adults collected in April.

DIAGNOSIS. A very distinctive small, brown-winged
Stigmella species characterized by the whitish
androconial scales on the upperside of the male
forewing, and by the androconial patch on the hindwing
(comprising black and whitish scales). See Remarks.

DISTRIBUTION. This appears to be a widespread
species in Belize, its habitats including disturbed rain-
forest and secondary forest areas close to settlements.

CONDITION OF TYPE MATERIAL. Wings not spread;
some specimens are poorly pinned, and a few lack
antennae.

MATERIAL EXAMINED.

Holotype 3, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, secondary forest close to re-
search station, 3—16.iv.1998 (Puplesis & Hill), genitalia
slide no. 29124 (BMNH).

Paratypes, 1d, data as holotype (BMNH); 96, 78,
San Ignacio, secondary forest at town edge, 17—
18.iv.1989 (Puplesis & Hill), genitalia slide no. 29126 @
(BMNH (14); VPU (2)); 1d, Pook’s Hill Nature Re-
serve (S of Teakettle Village), primary rain forest close
to main camp, 28—29.iv.1998 (Puplesis & Hill), geni-
talia slide no. AD0308 (VPU).

REMARKS. Some general features of the male genita-

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

lia (especially gnathos, transtilla and aedeagus) sug-
gest that S. pruinosa might be related to the S.
oxyacanthella-group (=S. crataegifoliella-group, sensu
Wilkinson & Scoble, 1979) which is very widely dis-
tributed in the Holarctic Region.

28. Stigmella ovata sp. n.
(Figs 35, 62, 148-151)

MALE (Fig.35). Forewing length: 2.3-2.5 mm. Wing-
span: 5.3—-5.6 mm. Head: palpi brown-grey to brown;
frontal tuft pale yellow to orange or pale ochre; collar
comprised of piliform cream scales; eye-caps cream to
white; antenna brown, ca. 31-32 segments. Thorax,
tegulae and base of forewing before first fascia grey-
brown to dark brown with some purple and blue-green
lustre (not always present). Antemedian fascia dis-
tinctly broadened towards dorsum, but generally more
slender than postmedian fascia, which tends to be
broader at costal margin or in middle; both fasciae
cream to pale yellow-cream, but postmedian usually
lighter and brighter, more intense. Area between fas-
ciae and apex of forewing dark brown with some
purple and blue-green lustre, often a little roughly
scaled (not smooth). Cilia grey-brown to dark brown;
no cilia-line. Underside of forewing brown. Hindwing
and cilia grey-brown; basal fifth of hindwing with row
of distinctly broadened scales on costal margin; these
are about one-third length of cilia but hardly visible as
they are concolorous with cilia. No androconia on
forewing. Legs grey-brown or dark brown. Abdomen
dark brown on upperside, grey-cream on underside; a
pair of short, closely set piliform anal tufts dark brown;
genital segments not contrasting in colour with ventral
surface of abdomen.

FEMALE. Unknown

GENITALIA ¢ (Figs 148-151). Capsule 429-446 um
long, markedly oval. Uncus broad at base, gradually
narrowing caudally, with two tiny, dark papillae at
caudal margin. Tegumen not extended into pseuduncus,
band-shaped. Gnathos inverted V-shaped; no central
plate; posterior process single, pointed caudally, lat-
eral arms long and slender. Valva 215—224 um long,
very broad in basal two-fifths, then abruptly narrowed;
apex very slender, pointed caudally. Transtilla with
slender transverse bar which is not fused at middle,
with papilla-like sublateral corners papilla-like, proc-
esses not developed. Vinculum very long and broad,
broadly rounded anteriorly, without emargination or
lateral lobes. Aedeagus ca. 232 um long, without apical
processes, characteristically broadened in caudal third.
Vesica with numerous small cornuti, most relatively
broad, and all pointing caudally. Juxta a short, broad
sclerite with a narrow bilobed caudal extension.

BIOLOGY. Adults collected in November—December.

39

DIAGNOSIS. This very distinctive species is easily
distinguished from all other Stigmella by the combina-
tion of huge, oval vinculum, inverted *V’-shaped

gnathos and posteriorly bilobed juxta. See Remarks.
DISTRIBUTION. Argentina.

CONDITION OF TYPE MATERIAL.
relatively well-preserved.

The type series is

MATERIAL EXAMINED.

Holotype 3, Argentina: Neuquen, Lago Lacar, Pucara,
26 20.xii.1978 (Mission Cientifica Danesa), genitalia
slide no. Diskus184 (ZMUC).

Paratypes, | d, dataas holotype; | d, data as holotype
but 28—29.xi.1981 (Nielsen & Karsholt), genitalia slide
no. Diskus185, forewing venation slide no. DiSkus 207
(ZMUC); 1d, Rio Negro, San Carlos de Bariloche,
Colonia Suiza, 800 m, 7.xii.1981 (Nielsen & Karsholt)
(ZMUC).

REMARKS. This species possesses many remarkably
archaic features: gnathos with a single process, primi-
tively shaped uncus, very large and rounded vinculum,
piliform collar and remarkably complete wing vena-
tion. In the forewing R, and R, are not fused and Cu is
surprisingly long; M, and M, are retained as separate
branches (Fig. 62) in contrast to other Stigmella (e.g.,
Fig. 63). This is the only species in which such a
character combination is known and we can offer no
explanation of affinities except to place ovata in the
genus Stigmella. This is an aberrant species in many
respects; among its atypical features are the structure
of the gnathos and uncus (Fig. 148) and vinculum
(Fig. 150); however, a few Palaearctic species do ex-
hibit such similar unusually-shaped sclerites.

29. Stigmella hylomaga (Meyrick, 1931)
(Fig. 36)

Nepticula hylomaga Meyrick, 1931: 415.
Stigmella hylomaga (Meyrick); Davis, 1984: 18.

ADULT (see Remarks) (Fig. 36). Forewing length:
2.7-2.8 mm. Wingspan: about 6.0-6.1 mm. Head: palpi
cream; frontal tuft very pale ochreous cream; collar not
very distinctive, brownish cream, comprised of lamel-
lar scales; eye-caps cream; antenna grey-brown with
weak purple lustre, about 37 segments. Thorax and
tegulae coppery brown with purple lustre. Basal part of
forewing of same colour as thorax, but apex and part of
basal fascia with much more intense purple lustre and
therefore appearing darker. Fascia oblique, distinctly
postmedial, whitish with some gold lustre.

BIOLOGY. The single specimen known was collected
in November.

DIAGNOSIS. May be distinguished from other
Neotropical species by its greater wingspan (about

40

6 mm) in combination with a broad, oblique, postme-
dian fascia surrounded by brown scales with a purple
lustre.

DISTRIBUTION. Argentina (Rio Negro region).

CONDITION OF TYPE MATERIAL. The holotype has no
hindwings or abdomen; the appendages of the head are
hardly visible and the left forewing is abraded.

MATERIAL EXAMINED.

Holotype | ex., Argentina: Rio Negro Territory, Lake
Correntoso, 15-25.xi.1926 (FW. & M. Edwards)
(BMNH).

REMARKS. This species was described by Meyrick
from a single specimen lacking hindwings and abdo-
men (mentioned in the original description). It is
impossible to ascertain the sex of this specimen. The
current placement of this species in Stigmella is provi-
sional, based on features such as the lamellar-scaled
collar and forewing venation, the latter examined in
situ rather than in a microscopic preparation.

30. Stigmella costalimai (Bourquin, 1962)
(Fig. 37)

Nepticula costalimai Bourquin, 1962: 31-32.
Stigmella costalimai (Bourquin); Davis; 1984: 18.

MALE. Unknown (see Remarks).

FEMALE (Fig. 37). Forewing length: about 2.0—
2.1mm. Wingspan: about 4.7-4.8 mm. Head: palpi
ochreous cream; frontal tuft brownish ochre. Collar
greyish, comprised of lamellar scales, lobes
well-separated from each other; eye-caps yellowish
cream; antenna brownish grey, number of segments
unknown. Thorax greyish, tegulae greyish but cream
posteriorly. Forewing glossy grey-bronze, but with
irregularly distributed, paler, brownish cream scales,
especially in distal half; most darker grey scales with
paler, brownish cream tips. Cilia greyish. Underside of
forewing brown. Hindwing and cilia brownish grey.
Abdomen fuscous.

GENITALIA 6,9. Unknown (see Remarks).

BIOLOGY. Adults reared in November.

DIAGNOSIS. Distinguishable among all other
Neotropical Stigmella and species from other genera
by the glossy grey-bronze forewing, with some irregu-
larly distributed paler scales.

DISTRIBUTION. Argentina (Buenos Aires region).

CONDITION OF TYPE MATERIAL. The single female
paratype examined lacks its abdomen; the wings are
detached and stored in a gelatine capsule. The holotype
and second paratype have not been examined.

R. PUPLESIS AND G.S. ROBINSON

MATERIAL EXAMINED.
Paratype, °, Argentina: Buenos Aires region, Tigre,
x.1957 (MACN).

REMARKS. S. costalimai was described from three
specimens in the Bourquin collection, Buenos Aires.
The original description of the species does not men-
tion the sex of the specimens of the type series. We
have been able to examine only a single female paratype
loaned by MACN through the kind offices of Dr. E.
Nieukerken (NNM).

31. Stigmella guittonae (Bourquin, 1962)
(Figs. 38, 217)

Nepticula guittonae Bourquin, 1962: 32-34.
Stigmella guittonae (Bourquin), Davis, 1984: 18.

MALE. Not examined (see Remarks).

FEMALE (Fig. 38). Forewing length: ca. 2.3 mm.
Wingspan: 4.9-5.0 mm. Head: palpi greyish cream;
frontal tuft ochreous yellow; collar indistinct, com-
prised of yellowish cream piliform scales; eye-caps
cream; antenna grey-brown, number of segments un-
known. Thorax and tegulae grey-brown. Forewing
grey-brown, indistinctly and very lightly irrorated with
darker brown scales, slightly darker than thorax; with
narrow, distinct, postmedian fascia formed by silver
lustrous (or white-metallic lustrous) scales; area bey-
ond fascia purple-brown; cilia grey-brown. Underside
of forewing fuscous. Hindwing relatively broad, brown-
ish grey; cilia concolorous. Legs grey-brown. Abdomen
dark grey.

GENITALIA 2 (Fig. 217). Total length 580-585 um.
Anal papillae weakly developed. Apophyses
posteriores slender and long (144-146 um). Apophy-
ses anteriores slightly longer and much broader.
Vestibulum slightly broadened, folded. Corpus bursae
small and narrow; no signa or pectinations visible.
Accessory sac relatively large, rounded, strongly
folded; ductus spermathecae sclerotized only proxi-
mally, and in a tiny area, at some distance from the
accessory sac.

BIOLOGY. Adults collected in February. According
to Bourquin (1962) the larvae are miners on Senecio
bonariensis (Compositae/Asteraceae) and Jussiaea
longifolia (Onagraceae); Brummitt (1992) treats
Jussiaea as a synonym of Ludwigia; the TROPICOS
database suggests Jussiaea longifolia may now be a
synonym of Ludwigia major (DC.) Ramamoorthy.

DIAGNOSIS. This species may be distinguished from
all other Neotropical nepticulids by the narrow and
distinctly postmedian oblique fascia formed by lus-
trous silver lustrous (or lustrous white-metallic) scales.

DISTRIBUTION. Argentina (Buenos Aires region).

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

CONDITION OF TYPE MATERIAL. The single female
paratype examined is unsatisfactorily pinned, rather
rubbed, with a damaged head and with both antennae
broken. On the data label the name is incorrectly
spelled as ‘guittoni’. The holotype and second paratype
have not been examined (see Remarks).

MATERIAL EXAMINED.
Paratype, °, Argentina: Buenos Aires Region, Tigre,
ii.1960, genitalia slide no. Diskus 2043 (MACN).

REMARKS. Wehave retained this species in Stigmella
as external examination of the paratype shows
Stigmella-type venation. S. guittonae was described
from three specimens in the Bourquin collection, Bue-
nos Aires. The original description of the species does
not mention the sex of the specimens of the type series.
We have been able to examine only a single female
paratype loaned by MACN through the kind offices of
Dr. E. Nieukerken (NNM).

ECTOEDEMIA Busck

32. Ectoedemia reneella Wilkinson, 1981
Ectoedemia reneella Wilkinson, 1981: 104, 105.
MALE (Described by Wilkinson, 1981: 104).

FEMALE. Unknown.

GENITALIA 6 (Described and figured by Wilkinson,
1981: 104, 105, fig. 6).

Adults collected in April—May.
USA (Florida).

BIOLOGY.

DISTRIBUTION.

33. Ectoedemia helenella Wilkinson, 1981
Ectoedemia helenella Wilkinson, 1981: 105-107.
MALE (Described by Wilkinson, 1981: 105, 106).
FEMALE. Unknown.

GENITALIA ¢ (Described and figured by Wilkinson,
1981: 106, fig. 7).

BIOLOGY. Adults collected in April.

DISTRIBUTION. USA (Florida).

34. Ectoedemia mesoloba Davis, 1978

Ectoedemia mesoloba Davis, 1978: 209-212.
Ectoedemia mesoloba Davis; Wilkinson & Newton,
1981: 85, 86.

MALE. Described and figured by Davis (1978: 209,
210, fig. 1) and by Wilkinson & Newton (1981: 85, 86,
fig. SO).

FEMALE. Unknown.

41

GENITALIA 6. Described and figured by Davis (1978:
210, 211, figs 15-17) and by Wilkinson & Newton
(1981: 85, 86, fig. 42).

BIOLOGY. Adults collected in November.

DISTRIBUTION. USA (extreme northwestern Florida).

35. Ectoedemia species 29105
(Figs 40, 152, 153)

MALE (Fig. 40). Forewing length: 3.2 mm. Wing-
span: 7.1mm. Head: palpi cream; frontal tuft
brown-black, very large and dark; collar blackish (not
off-white as in Nearctic reneella Wilkinson), com-
prised of piliform scales; eye-caps cream, large; antenna
brown-grey (not off-white as in reneella), about 56
segments. Thorax and tegulae greyish yellow with
dark brown and blackish scales especially abundant
anteriorly on tegulae. Forewing greyish yellow, irregu-
larly but not densely irrorated with blackish brown
scales; a few dark scales on cilia also. Cilia greyish
yellow distally, grey tornally. Underside of forewing
brown with green-blue lustre. Hindwing relatively
broad, distinctly narrowed apically, cream or greyish
cream (at certain angles); hair-pencil of long piliform
scales along costa of forewing base well-developed,
cream and surrounded with tiny blackish scales. Cilia
of hindwing cream to greyish cream. Legs yellowish
cream to ochreous cream, shaded with black at front.
Colour of abdomen unknown.

FEMALE. Unknown.

MALE GENITALIA (Figs 152, 153). Capsule 490 pm
long. Uncus absent. Pseuduncus short, broad, band-
shaped, with laterodorsal setose papillae. Tegumen
short. Gnathos W-shaped, with relatively short but
broad caudal process; lateral arms long and
well-sclerotized; central plate undeveloped. Valva
377 um long, relatively slender, gradually narrowed
towards apex, slightly curved inwards, with short broad
papillated median extension on dorsal surface.
Transtilla with long and slender transverse bar and
long slender sublateral processes. Ventral plate of vin-
culum short, with triangular lateral lobes; anterior
emargination relatively shallow and broad. Aedeagus
547-554 um long, with two pairs of carinae: lateral
carinae short, but well-sclerotized; dorsal carinae lob-
ate, spine-like at apices; ventral carinae hardly
developed, represented by a short single plate with
triangular emargination from caudal margin. Vesica
with three comb-like sclerotized plates each bearing
5-6 spines, a few large spinose cornuti and numerous
scattered tiny cornuti.

BIOLOGY. Adults collected in April. Larvae may be
bark miners.

DIAGNOSIS. Belongs to the highly specialized

42

subgenus Zimmermannia Hering, which is character-
ized by pronounced costal emargination of the male
hindwing, large ventral carinae on the aedeagus, elon-
gated valvae, extremely long and narrow bursa in the
female genitalia, and patternless forewing. Larvae of
Holarctic representatives this subgenus are bark-miners
with an atypically high number of instars — six to eight.

This species most resembles to E. reneella
Wilkinson, from the USA (Florida). However, the
valva is narrower basally, the pseuduncus broader, the
lateral carinae more elaborated, the dorsal carinae
abruptly narrowed the apex into spine-like processes,
the ventral carinae weakly developed and demarcated,
the transverse bar of the transtilla fused in the middle.
The antennae are brown and the collar is black, not
white as in reneella.

The description of Ectoedemia species 29105 repre-
sents the first and only record of the subgenus
Zimmermannia in the rainforest of the Neotropical
Region; there are no other species currently known
south of Mexico.

DISTRIBUTION. Belize.

CONDITION OF MATERIAL.
able is well-preserved.

The single specimen avail-

MATERIAL EXAMINED.

1d, Belize: Cayo District, Chiquibul Forest Reserve,
Las Cuevas, 3-16.iv.1998 (Puplesis & Hill), genitalia
slide no. 29105 (BMNH).

REMARKS. This interesting species has been described
but not named, because we are not persuaded from the
single known specimen that this species is more than a
distinct geographical variant of reneella Wilkinson, to
which it is undoubtedly closely related.

36. Ectoedemia fuscivittata sp. n.

(Figs 39, 154-156)

MALE (Fig. 39). Forewing length: 1.7-1.8 mm. Wing-
span: 4.1-4.3 mm. Head: palpi cream to ochreous
cream; frontal tuft pale ochre to pale orange-ochre;
collar indistinct, pale ochre, comprised of piliform
scales; eye-caps ochreous cream, moderately large;
antenna brownish or greyish to pale brownish ochre,
42-44 segments. Thorax anteriorly ochreous yellow,
elsewhere metallic grey; tegulae ochreous yellow with
a few greyish scales anteriorly. Forewing up to fascia
(in basal 3/5) metallic grey with distinctive greenish
blue and some indistinct purplish lustre; narrow area
along costa, particularly before fascia and on tornus, as
well as area beyond fascia with ochreous cream scales;
fascia distinctly postmedian, oblique, fuscous brown;
some specimens with dark brown scales in the area
before cilia. Cilia metallic greyish. Underside of

R. PUPLESIS AND G.S. ROBINSON

forewing grey-brown. Hindwing and cilia greyish to
brownish grey. A long patch of whitish scales may be
visible on basal half of hindwing upperside, but these
androconia may not always be distinctive. Legs cream
shaded with grey-fuscous laterally. Abdomen fuscous
brown on upperside, brown or brownish on underside;
genital segments mostly covered by dark (predomi-
nantly brown) scales, not contrasting with main colour
of abdomen.

FEMALE. Unknown.

GENITALIA 6 (Figs 154-156). Capsule 268-270 um
long. Pseuduncus small, rounded, distinctly papillated.
Dorsal plate of tegumen simple, small. Gnathos with
triangular caudal process, slender lateral arms, and
small rounded central plate. Valva 150-152 um long,
triangular, with more or less straight inner margin,
tapering into a pointed, subcaudally-directed apical
process. Basal margin of valva strongly sclerotized.
Transtilla without transverse bar, with long and slender
sublateral processes. Juxta absent, valvae connected
by membranous basal. Vinculum very small, with com-
pact, distinctively-shaped and well-sclerotized lateral
lobes; anterior emargination of vinculum broad but
shallow, almost quadrate. Aedeagus 23 1—235 um long,
strongly bulged in basal one-half to one-third, without
carinae; vesica with only very few tiny and indistinct
cornuti.

BIOLOGY. Adults collected in April.

DIAGNOSIS. Belongs to the nominal subgenus
Ectoedemia Busck, which has a wide distribution, but
is here recorded for the first time from the Neotropical
Region south of Mexico. Although many Ectoedemia
species are difficult to separate, fuscivittata is easily
recognizable: a combination of features such as the
subcaudally-directed valval process, basally broad-
ened aedeagus, very short vinculum, and dark oblique
forewing fascia make this species distinctive. The ab-
sence of the transverse transtilla bar in this species is
the only known occurrence of this character in the
genus.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype and
one paratype are well-pinned and well-preserved (but
unspread) specimens; the second paratype is unsatis-
factorily pinned.

MATERIAL EXAMINED.
Holotype 6, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slides no. 29107 (BMNH).

Paratypes, 2d, data as holotype, genitalia slide no.
AD 0302 (BMNH, VPU).

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE
FOMORIA Beirne

37. Fomoria molybditis (Zeller, 1877) comb. n.
(Figs 41, 157, 158)

Nepticula molybditis Zeller, 1877: 455-456.
Stigmella molybditis (Zeller); Davis, 1984: 18.

MALE (Fig. 41). Forewing length: about 4.0 mm.
Wingspan: 8.5—8.6 mm. Head: palpi pale grey-brown;
frontal tuft pale ochre; collar pale ochre, rather indis-
tinct, comprised of piliform scales; eye-caps ochreous
cream; antenna brownish, number of segments un-
known (antennae broken in single specimen available).
Thorax and tegulae dark brown, lustrous. Forewing
without pattern, brown and lustrous; costal margin and
apical area darkened by numerous deep brown scales
which are scattered universally across forewing but
concentrated along costa and at apex; distinctly edged
by dark brown scales just before cilia. Cilia grey-
brown. Underside of forewing brown. Hindwing
lanceolate, rather broad, grey-brown, cilia brown. No
androconia on forewing or hindwing. Legs brown.
Colour of abdomen unknown.

FEMALE. Unknown.

GENITALIA 6 (Figs 157, 158). Capsule 560 um long.
Pseuduncus with two well-separated lateral lobes.
Uncus inverted V-shaped. Tegumen small. Gnathos
with slender posterior process and almost equally de-
veloped but longer lateral arms. Valva 235 um long;
apical half very slender and turned inwards; basal half
broad and with small inner lobe; long and very slender
spine-like process on dorsal side of valva, reaching
gnathos caudally, fused basally with transtilla.
Transtilla with very large transverse bar that is
well-sclerotized only laterally; without sublateral proc-
esses. Vinculum narrow, elongate, longer than valva;
anterior emargination shallow. Aedeagus 544 um long,
with two (sclerotized and weakly sclerotized) lobe-like
apical processes, one sclerotized, the other weakly so.
Vesica with numerous spine-like cornuti mostly
arranged in a long row, but with separate more scat-
tered cornuti in basal half.

BIOLOGY. Adults collected in February.

DIAGNOSIS. This species may be distinguished from
other Fomoria and Neotropical nepticulids by the unu-
sually long transtilla plate in combination with the
long and sinuous basal process of the valva and long
narrowed vinculum. It most resembles, and is most
closely related to diskusi but is easily separable by
external and male genital features (see Diagnosis for
diskusi).

DISTRIBUTION. Colombia.

43

CONDITION OF TYPE MATERIAL. The single specimen
is in good condition, but both antennae are missing.

MATERIAL EXAMINED.

Holotype 3, Colombia: nr. Bogota, foot of Guadalupe
[‘Fusse des Guadalupe bei Bogota’], 11.11.1871
(Nolcken), genitalia slide no. 25651 (BMNH).

REMARKS. ‘The present position of this species within
Fomoria is not certain and may be clarified when more
and reared material is available. In order to preserve
the unique and fragile holotype, the wing venation of
this species has not been studied by making a wing
preparation. Little can be deduced from external exam-
ination, but even details of venation might not be
particularly helpful as reduction of veins occurs re-
peatedly and independently throughout the
Nepticulidae. The morphology of the male genitalia of
molybditis is very compatible with that of a Fomoria
(see Introduction), and precludes its inclusion in
Stigmella (=Nepticula) where it was originally placed.
The species slightly resembles also some Parafomoria.
However, the latter genus (currently known mainly
from the Mediterranean region, feeding on Cistaceae)
is characterized by the lateral arms of the vinculum
being expanded and often wrinkled, unlike those of
molybditis, and by the lack of vein R, , in the forewing.
In the possibly related F: diskusi the cell is closed
(unusual in Fomoria) but R,,, is present, in contrast to
Parafomoria. On balance, placement of molybditis in
Fomoria rather than Parafomoria is appropriate.

38. Fomoria diskusi sp. n.
(Figs 42, 64, 159, 160, 219)

MALE (Fig.42). Forewing length: 1.9—2.3 mm. Wing-
span: 4.8-5.3 mm. Head: palpi cream to ochreous
cream; frontal tuft orange-ochre, occasionally brown
at front; collar pale ochreous, indistinct, comprised of
piliform scales; eye-caps cream; antenna pale brown-
ish ochre, ca. 44 segments. Thorax ochreous cream
anteriorly, fuscous brown posteriorly; tegulae ochre-
ous cream, occasionally with few fuscous brown scales.
Forewing fuscous brown with broad ochreous cream
area along costa connecting with broad oblique post-
median ochreous cream fascia; some fuscous brown
scales sparsely distributed on pale markings of
forewing. Cilia ochreous cream. Underside of forewing
brown to dark grey-brown. Hindwing and cilia grey-
cream to greyish. No androconia on hindwing or
forewing. Legs ochreous cream shaded with
grey-brown on front. Abdomen grey-brown or fuscous
on upperside, ochreous cream on underside; anal tufts
ochreous cream.

FEMALE. Very similar to male, but sometimes paler,
especially on underside of forewing. Antenna ca. 38
segments.

44

GENITALIA 6 (Figs 159, 160). Capsule 434 um long.
Pseuduncus distinctly bilobed with caudal setae. Un-
cus inverted V-shaped. Gnathos inverted V-shaped,
with relatively short caudal element, well-sclerotized
lateral arms, without central plate. Valva 230-232 um,
slender in apical two-fifths, with apical spine-like proc-
ess; basally with broad ventral lobe and two huge
elongate caudally-directed processes. Transtilla with-
out sublateral processes, a very long but narrow plate,
slightly broadened anteriorly and strongly sclerotized
in middle. Vinculum very long, with shallow semicir-
cular anterior emargination and small triangular lateral
lobes. Juxta absent. Aedeagus 378 um, relatively slen-
der, especially in basal third, simple, with two pointed
apical carinae. Cornuti absent.

GENITALIA ¢(Fig.219). Total length 560-563 um. S8
and T8 broadly rounded. Apophyses posteriores about
154 um long, unusually broad for most of their length,
abruptly and strongly narrowed at apices. Apophyses
anteriores more slender than apophyses posteriores,
approximately same length. Ductus with complex
bulged lobate sclerotization. Corpus bursae small, oval,
with one rather distinct spine-like sclerite, but without
any visible signa. Accessory sac small, rounded; duc-
tus spermathecae broad, strongly sclerotized and with
7-8 convolutions; length of ductus spermathecae ex-
ceeds that of corpus bursae.

BIOLOGY. Adults collected in April.

DIAGNOSIS. An outstandingly distinctive species
among Fomoria and all others nepticulids by dint of its
unique forewing pattern (Fig. 42); in the resting posi-
tion resembles a seed surrounded with white. The
distinctive male genitalia differ from those of other
species by the double basal processes of the valva
together with the isolated valval lobe, unusual transtilla
plate, and by the absence of cornuti in the long and
slender aedeagus. F: diskusi is somewhat similar to and
possibly closely related to molybditis from which it
clearly differs in all the characters listed above, except
the form of the transtilla.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype and
most paratypes well-preserved; the single female
paratype is slightly rubbed and the detached hindwings
have been pinned in a gelatine capsule.

MATERIAL EXAMINED.
Holotype 6, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 28844 (BMNH).

Paratypes, 11d, 1 2, data as holotype, genitalia slide
nos 28845, 291306, 28846 2 (BMNH; VPU).

REMARKS. This remarkable species is named for
Arunas Diskus (VPU), to whom we are extremely

R. PUPLESIS AND G.S. ROBINSON

grateful for his help in dissecting many specimens of
Neotropical nepticulids and for his support for our
work on leaf-miners.

This species is placed in Fomoria because the mor-
phology of the male genitalia matches well with the
current Fomoria genus concept; the unusually reduced
wing venation (Fig. 64) lends additional support. The
vaginal lobe-like sclerotizations and short bursa
resemble those in the genus Parafomoria; however, in
the male genitalia the vinculum arms are not expanded
or wrinkled, and forewing vein R, , is present (lacking
in Parafomoria).

39. Fomoria species 29122
(Figs 44, 161-163)

MALE (Fig. 44). Forewing length: ca. 1.8 mm. Wing-
span: ca. 4.2 mm. Head: palpi cream; frontal tuft pale
ochre; collar ill-defined, cream; eye-caps cream; an-
tenna unknown. Thorax and tegulae blackish. Forewing
brownish black with gold and weak purple lustre; a few
glossy silver markings on forewing: at base of forewing,
a narrow oblique median fascia and two narrow ter-
minal spots which tend to fuse forming a terminal
fascia. Cilia grey; some long lamellar wing scales
overlap cilia. Underside of forewing creamy brown.
Hindwing lanceolate, brownish cream, cilia brownish.
No androconia on forewing or hindwing. Abdomen
ochre-brown on upperside, ochreous cream on under-
side; genital segments (mainly valvae) ochreous cream,
easily visible, but not contrasting in colour with ventral
surface of abdomen.

FEMALE. Unknown.

GENITALIA ¢ (Figs 161-163). Capsule 340 um long.
Pseuduncus deeply bilobed, lobes large, triangular.
Uncus inverted V-shaped, with short and blunt caudal
process. Gnathos with single caudally directed process
and equally developed lateral arms. Valva 120 um
long, very deeply divided into a longer, distally inturned
dorsal lobe and triangular ventral lobe. Transtilla with
large lobate transverse bar with remarkably extended
and pointed sublateral angles. Vinculum very long and
very broad, almost without anterior emargination.
Aedeagus 160-166 um long, with numerous large
spine-like cornuti on vesica; no apical carinae are
visible in the single specimen available.

BIOLOGY. Adults collected in April.

DIAGNOSIS. Easily distinguished from other Fomoria
and all other nepticulids by the unique shape of the
transtilla (Fig. 161), and by the long vinculum in
combination with the divided valva with triangular
ventral lobe and narrow dorsal lobe.

DISTRIBUTION. Belize.

CONDITION OF MATERIAL. The single available speci-

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

men is in very poor condition: the body is destroyed,
the head is glued on a pin separately, and the wings are
abraded.

MATERIAL EXAMINED.

1d, Belize: Cayo District, Chiquibul Forest Reserve,
Las Cuevas, 3—16.iv.1998 (Puplesis & Hill), genitalia
slide no. 29122 (BMNH).

REMARKS. This very remarkable and undoubtedly
new species is described but it is not named because of
the very poor condition of the single known specimen.

40. Fomoria latipennata sp. n.
(Figs 43, 164, 165)

MALE (Fig.43). Forewing length: 2.4—2.5 mm. Wing-
span: 5.4-5.5 mm. Head: palpi cream; frontal tuft
brownish ochre, rather dark; collar cream, comprised
of piliform scales, indistinct; eye-caps large, ochreous
cream, distally shaded with greyish; antenna grey-
brown, about 26 segments. Thorax and tegulae grey or
dark grey. Forewing broad; basal half grey with weak
bronze gloss; apical half of forewing fuscous, the
demarcation from the basal half oblique; proximal
third of apical fascia and apex covered with blackish
brown scales with some yellowish lustre; region
between these blackish areas dark grey with metallic
gloss. Cilia grey with some shorter intermixed fuscous
scales. Underside of forewing grey brown. Hindwing
relatively broad, ochreous brown, cilia grey-brown.
No androconia on hindwing or forewing. Legs irregu-
larly coloured, brownish cream to predominantly
grey-brown. Colour of abdomen unknown.

FEMALE. Undescribed. A single female specimen is
tentatively associated with the male but is not described
as its status is conjectural (see Remarks).

GENITALIA 6 (Figs 164, 165). Capsule 415 pum.
Pseuduncus with two large, smoothly rounded
sublateral lobes and one huge central lobe. Uncus
smoothly sclerotized, caudally bilobed, with two pairs
of large setae on ventral side. Dorsal plate of tegumen
moderately large, simple. Gnathos with large rounded
caudal process, long slender lateral arms, and weakly
developed (anteriorly broadly rounded) central plate.
Valva 256 um long, broadened at base, with spine-like
median process and long and very slender apical proc-
ess from apex of broad valval lobe. Transtilla with
narrow and elongate transverse bar and very long
sublateral processes extending beyond vinculum. Juxta
absent. Vinculum large, ventral plate trapezoidal,
gradually narrowed anteriorly, without lateral lobes
and anterior emargination. Aedeagus 256 pum long,
swollen in apical half and weakly sclerotized at basal
margin; with three very long horn-like apical cornuti
and one moderately large spinose central cornutus.
The longest cornutus is strongly curved outwards.

45
GENITALIA 2 (see Remarks).

BIOLOGY. Adults collected in April.

DIAGNOSIS. The distinctive structure of the male geni-
talia, particularly the bilobed uncus, trilobed
pseuduncus, long sublateral processes of the transtilla,
and the fuscous-banded forewing distinguish this

species from all other nepticulids.
DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL.
well-preserved.

The holotype is

MATERIAL EXAMINED.
Holotype 3, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 28968 (BMNH).

Excluded from paratype series: | 9, dataas holotype,
genitalia slide no. 28969 (BMNH) (see Remarks).

REMARKS. We have very tentatively associated a fe-
male with this male but have not (above) allotted it
paratype status. The genitalia (slide no. 28969, BMNH)
exhibit a narrowed abdominal tip, very long and broad
posterior and anterior apophyses, folded vestibulum,
relatively short oval corpus bursae (without signa or
pectinations), rather small accessory sac and short,
distally strongly sclerotized ductus spermathecae (with
2.5—3 convolutions). External features of this female
differ somewhat from those of the male holotype; the
forewing is more roughly scaled and has a distinct
metallic lustre and golden gloss.

ACALYPTRIS Meyrick

41. Acalyptris bovicorneus sp. n.
(Figs 45, 65, 166-169, 220)

MALE (Fig.45). Forewing length 2.3—2.5 mm. Wing-
span: 5.4—5.8 mm. Head: palpi cream; frontal tuft pale
to dark orange-ochre; collar comprised of cream
piliform scales; eye-caps cream; antenna brown to
fuscous, ca. 42-46 short segments. Thorax and tegulae
varying from grey-brown and slightly irrorated paler,
to fuscous, almost black, without paler irroration.
Forewing colour varying from greyish cream irrorated
with densely scattered brown-grey or fuscous scales to
completely fuscous or black with no pale scales; a tiny
greyish cream tornal spot, sometimes indistinct and
fused with cilia of same colour, but always present.
Cilia cream distally, greyish cream on tornus; blackish
or dark grey scales overlap on cilia and may sometimes
form an indistinct cilia line. Underside of forewing
fuscous. Hindwing lanceolate, brownish or greyish;
cilia greyish or cream. No androconia on hindwing or
forewing. Legs cream with grey to fuscous irregular
lateral shading. Abdomen dark brown, on underside

46

sometimes paler due to cream scales; scaling of genital
segments ochreous cream, strongly contrasting in col-
our with remainder of abdomen.

FEMALE. Similar to male.

GENITALIA 6 (Figs 166-169). Capsule 456-464 um
long. Tegumen extended into triangular, caudally pap-
illated and setose pseuduncus. Uncus complex, with
semicircular transverse bar with tiny lateral spinose
processes close to gnathos arms, small and shallow
ventrally-directed central lobe and two slender
sublaterally-directed caudal processes. Gnathos with
short triangular caudal process, short but broad and
strongly sclerotized lateral arms, and small anteriorly-
extended central plate. Tegumen forming lateral
rod-like sclerites lying almost along valvae. Valva
310-315 pm long, simple, gradually narrowed towards
apex and with distinctive brush of broad setae in apical
quarter forming a pectinifer. Transtilla absent, 1.e., no
transverse bar; sublateral processes slender and rela-
tively long. Posterior margin of vinculum with tiny,
spine-like sclerite that may represent a fused vestige of
the juxta (but see below). Ventral plate of vinculum
very short, but with large triangular and anteriorly
rounded lateral lobes; anterior emargination deep, in-
verted V-shaped. Aedeagus 437-482 ym (including
long processes); tube narrow, with two very long
(maximally about 210 um) asymmetrical apical cari-
nae, two (sometimes three) small spinose dorsal carinae
and one large spinose ventral carina; ventral carina
connected with aedeagus tube via long transverse ex-
tension that forms a virtual bridge between the valvae
and may represent the juxta or part of the juxta; all
carinae connected by a strong quadrate sclerotization.
Vesica with very tiny spinose cornuti.

GENITALIA ¢(Fig. 220). Total length 995-1024 um.
Abdominal tip almost square. Anal papillae hardly
developed, ill-defined, broadly rounded. Apophyses
anteriores broad and long, terminating anteriorly in a
slender inward-curved claw-shaped process. Apophy-
ses posteriores very slender, straight, distinctly longer
than apophyses anteriores. Vestibulum with complex
oval sclerotization. Corpus bursae irregularly shaped,
extended anteriorly and laterally into an oblique sac.
Signa on both sides of bursa: tiny pectens forming a
chain of closed cells in two parallel bands. Accessory
sac oval, relatively small; ductus spermathecae short,
with 2.5 small convolutions close to accessory sac.

BIOLOGY. Adults collected in April in both rainforest
and disturbed secondary forest close to settlements.

DIAGNOSIS. Differs external from many Neotropical
Acalyptris in having a monotonous forewing pattern,
but as such is similar to platygnathos, species 29135,
unicornis, fortis, bifidus, tenuijuxtus and aberrant speci-
mens of other species. The male genitalia differ

R. PUPLESIS AND G.S. ROBINSON

spectacularly from those of all other Neotropical species
in the presence of a valval pectinifer; bovicorneus is
also easily distinguishable by the two very long and
divergent aedeagal carinae, or by the two-pronged
uncus in combination with the lack of a transtilla.
The valval pectinifer differentiates bovicorneus from
Acalyptris from other parts of the world except for
some species of the Asiatic repeteki-group; however,
none of these exhibit either a two-pronged uncus, very
long lateral aedeagal carinae, a short triangular poste-
rior gnathos process, or a similarly shaped pseuduncus.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype is
well-preserved, but one paratype has its abdomen miss-
ing, while another has abraded forewings.

MATERIAL EXAMINED.
Holotype 6d, Belize: Cayo District, San Ignacio, dis-
turbed secondary forest, 17—18.iv.1998 (Puplesis &
Hill), genitalia slide no. 29115 (BMNH).

Paratypes: 3d, 12, Chiquibul Forest Reserve, Las
Cuevas, 3—16.1v.1998 (Puplesis & Hill), genitalia slide
nos 29116 2, AD0307d (BMNH (3); VPU (1)).

REMARKS. This is an apparently very isolated, re-
markable Neotropical species; its modest external
appearance belies a bizarre male genital morphology
with numerous highly specialized features. The pres-
ence of a pectinifer may well, however, be a
plesiomorphy.

42. Acalyptris martinheringi sp. n.
(Figs 46, 170, 171)

MALE (Fig. 46). Forewing length: 2.1—2.5 mm. Wing-
span: 4.7—5.6 mm. Head: palpi cream; frontal tuft large,
ochreous brown or brown; collar comprised of cream
piliform scales; eye-caps large, cream; antenna brown
or ochreous brown, 41-43 short segments. Thorax
cream, usually with some ochre or brownish scales;
tegulae cream intensely shaded with brown. Forewing
cream irrorated with ochreous, ochreous brown and
dark brown; dorsal area of forewing base predomi-
nantly cream, weakly irrorated; very narrow area of
forewing base along costa densely shaded with small
brown or fuscous brown scales; apex pale; irregular
dark spots scattered everywhere on forewing, espe-
cially along tornus and costal margin. Cilia cream with
some brown to fuscous brown scales. Underside of
forewing brownish except for small elongated basal
areas where it may be cream. Hindwing lanceolate,
ochreous cream, cilia cream. No androconial patches
on hindwing or forewing. Legs cream shaded laterally
with brown. Abdomen ochreous brown on upperside,
brownish cream or cream on underside.

FEMALE. Unknown.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

GENITALIA 6 (Figs 170, 171). Capsule 402-418 pm
long. Tegumen extended into large broadly rounded
and setose pseuduncus. Uncus inverted V-shaped, with
tiny caudal process. Gnathos with large caudal proc-
ess, broad and strongly sclerotized lateral arms; central
plate, as such, undeveloped. Tegumen with lateral
rod-like sclerites along capsule dorsal to valvae. Valva
261-273 um long, narrow, without lobes or processes,
strongly narrowed in basal third. Transtilla with trans-
verse bar and long sublateral processes. Juxta strongly
sclerotized, elaborated, with two sharp, outwardly
curved, horn-like processes from plate-like extension
of vinculum. Vinculum very short and broad, with
broad triangular lateral lobes; anterior emargination
broad and shallow. Aedeagus 378-385 um long, slen-
der, with two large horn-like, curved lateral carinae,
one of them larger than the other. Vesica with numer-
ous spine-like cornuti, most of them very small.

BIOLOGY. Adults collected in April in rainforest and
in disturbed secondary forest habitats close to settle-
ments.

DIAGNOSIS. Among congeneric Neotropical species,
martinheringi resembles externally fortis, bifidus,
species 29135, unicornis, platygnathos, species 29140,
tenuijuxtus, bicornutus and aberrant specimens of other
species in having a lightly irrorated forewing. How-
ever it may be distinguished in male genital structure
by the large and strongly developed juxta with horned
lateral extensions; additionally it can be characterized
by the basally narrowed valva and broad pseuduncus.
It differs from all other Acalyptris species in the highly
developed and specialized two-branched juxta with
lateral outgrowths.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype and
most paratypes are well preserved, but the forewings
are not spread.

MATERIAL EXAMINED.

Holotype d, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.1v.1998 (Puplesis & Hill),
genitalia slide no. 29109 (BMNH).

Paratypes, 3d, data as holotype, genitalia slide nos
29129, AD0304 (BMNH (2); VPU (1)); 16, San
Ignacio, secondary forest, 17—18.iv.1998 (Puplesis &
Hill) (BMNH#).

REMARKS. This remarkable species is named in hon-
our of Prof. Martin Hering, a pioneer of studies of
leaf-mining moths and their biology, whose Memorial
Fund in part financially supported this project. This is
a phylogenetically isolated species with unique juxta
structure but conservative and plesiomorphic
pseuduncus and uncus.

47
43. Acalyptris fortis sp. n.
(Figs 47, 174-177)

MALE (Fig.47). Forewing length: 1.8—1.9 mm. Wing-
span: 4.1—4.3 mm. Head: palpi pale brownish cream;
frontal tuft orange-ochre; collar comprised of ochre-
ous piliform scales; eye-caps cream with a very few
brownish scales; antenna brown, ca. 33-34 segments.
Thorax predominantly cream, slightly irrorated with
brown. Forewing fuscous, distally cream; cream back-
ground of forewing irrorated with very numerous
fuscous, fuscous-brown and brownish scales; narrow
basal area along costa almost black; dark scales over-
lap onto cream cilia. Underside of forewing
grey-brown. Hindwing narrow, lanceolate, ochreous
cream; cilia cream. No androconial patches on
hindwing or forewing. Legs cream with irregular fus-
cous and grey lateral shading. Abdomen cream.

FEMALE. Unknown.

GENITALIA 6 (Figs 174-177). Capsule 330-342 um.
Tegumen caudally extended caudally into a large,
caudally rounded and setose pseuduncus. Uncus in-
verted V-shaped, with short, slender caudal process.
Gnathos with long, slender caudal process and lateral
arms; central plate small, very broadly rounded
anteriorly. Vinculum with lateral rod-like sclerites un-
derlying valvae. Valva 243-268 um, abruptly narrowed
in apical third and broad in basal two-thirds, with huge
bifid inward-directed spine-like process. Transtilla with
long slender transverse bar and slender sublateral proc-
esses. Ventral plate of vinculum very short and broad,
with broadly triangular lateral lobes; anterior
emargination shallow. Aedeagus 378-390 um long,
simple at base but very complex in swollen apical
region, with two huge horn-like lateral carinae and one
horn-like carina in between them; apex with one or two
rounded lobes, which at a certain angle may appear as
spine-like sclerotization. Vesica with numerous tiny
cornuti and three spine-like cornuti, two very large and
one that is half the size placed apart from them.

BIOLOGY. Adults collected in April in rainforest, and
in disturbed secondary forest close to settlements.

DIAGNOSIS. Among congeneric Neotropical species,
fortis resembles externally martinheringi, bifidus,
species 29135, unicornis, platygnathos, species 29140,
tenuijuxtus, bicornutus and aberrant specimens of other
species in having an irrorated forewing. However, in
the male genitalia it has a sharp, strong and bifurcated
process on the inner side of the valva which is a unique
feature; additionally it may be easily separated by the
long, strongly developed aedeagal carinae of aedeagus
in combination with the very broad pseuduncus,
broader than in any other Neotropical species. It may

48

be distinguished from all other Acalyptris by the valval
process or by the combination of very broad
pseuduncus and very long aedeagal carinae.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype speci-
men is abraded; the paratype specimens (except one)
are in poor condition and stored in gelatine capsules.

MATERIAL EXAMINED.

Holotype 3, Belize: Cayo District, San Ignacio, sec-
ondary forest, 17—18.iv.1998 (Puplesis & Hill),
genitalia slide no. 29131 (BMNH).

Paratypes: 1d, data as holotype, genitalia slide no.
29136 (BMNH); 1d, Chiquibul Forest Reserve, Las
Cuevas, 3-16.iv.1998 (Puplesis & Hill), genitalia slide
no. 29139 (BMNH); 26, Pook’s Hill Nature Reserve,
S. of Teakettle Village, 28-29.iv.1998 (Puplesis &
Hill), genitalia slide nos AD 0311 d (BMNH; VPU).

REMARKS. A very specialized species with strongly
‘armed’ and relatively large male genitalia, hence the
specific name.

44. Acalyptris hispidus sp. n.
(Figs 48, 172, 173)

MALE (Fig. 48). Forewing length: 2.6 mm. Wing-
span: 5.8 mm. Head: palpi ochreous cream or cream;
frontal tuft intense orange-ochre; collar cream, com-
prised of piliform scales; eye-caps large, orange-cream;
antenna ochreous brown, about 48 segments. Thorax
and tegulae grey. Forewing ground colour predomi-
nantly cream with sparsely scattered grey-brown scales;
dark markings comprise a basal area elongated along
costa, one or two small median spots and a strongly
oblique apical fascia (or anastomosing spots); dark
markings predominantly blackish brown, at certain
angles appearing almost black; some fuscous scales on
cilia also; anal area of forewing cream, densely covered
with pale grey-brown scales; oblique area between
median spots and apical fuscous fascia (or anastomos-
ing spots if fascia not developed) of ground colour; tint
of grey-brown scales varying with angle of view, at
certain angles they may look more cream than grey;
cilia grey. Underside of forewing fuscous brown.
Hindwing grey; in basal two-fifths with parallel elon-
gate stripes of fuscous grey scales along costa and in
middle; scales between these longitudinal stripes
cream. Legs cream with fuscous frontal and lateral
shading.

Unknown.

GENITALIA 6 (Figs 172, 173). Capsule 395-398 um.
Pseuduncus distinctly papillated, relatively long and
narrow, rounded caudally. Uncus broad inverted V-
shaped, arms swollen laterally. Tegumen with small
dorsal plate and long rod-like lateral projections along

FEMALE.

R. PUPLESIS AND G.S. ROBINSON

genital capsule. Gnathos with large caudal process,
broad lateral arms and quadrate central plate extended
and strongly sclerotized anteriorly. Valva 220 um,
narrow and simple, without processes or lobes.
Transtilla (broken in holotype, Fig. 172) with trans-
verse bar. Juxta absent. Ventral plate of vinculum
relatively small, with distinct triangular (anteriorly
almost pointed) lateral lobes; anterior emargination
semicircular. Aedeagus 309 pum, relatively broad, esp-
ecially in apical third, at its widest approximately
twice width of valval base; with one rounded apical
carina. Vesica in apical part of aedeagus with group of
large and broad-based cornuti together with numerous
scattered tiny cornuti and with large medial plate-like
sclerotization.

BIOLOGY. Adults collected in April.

DIAGNOSIS. This species externally resembles only
laxibasis and novenarius among Neotropical
Acalyptris, but it differs from both in the presence of
androconia on the hindwing and the significantly larger
wingspan. In the male genitalia it differs from all
Neotropical Acalyptris species, including laxibasis and
novenarius, in the combination of specialized gnathos
with broad quadrate basal extension and narrow, very
simple and unarmed valva. From Acalyptris species
elsewhere with a similar valva, it differs in the charac-
teristic structure of the gnathos and in the presence of
a forewing postmedian fascia.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype is
well-preserved; in the genitalia preparation the right
valva has been removed and mounted separately, a
procedure that unfortunately destroyed part of the
transtilla.

MATERIAL EXAMINED.

Holotype 3d, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3-16.iv.1998 (Puplesis & Hill),
genitalia slide no. 29104 (BMNH).

REMARKS. This species exhibits a surprisingly con-
servative male genital structure, with the exception of
the specialized gnathos. In contrast, the strongly devel-
oped forewing pattern and hindwing androconia are
apomorphic within the context of Acalyptris.

45. Acalyptris novenarius sp. n.
(Figs 49, 178, 179)

MALE. (Fig. 49). Forewing length: 2.2—2.3 mm.
Wingspan: 4.8—5.1 mm. Head: palpi cream to greyish
or brownish cream; frontal tuft orange-ochre to pale
ochre; collar sometimes indistinct, pale ochre or ochre-
ous cream, comprised of piliform scales; eye-caps
cream, usually with some greyish scales distally; ant-
enna ochreous brown or pale brown, about 56 short

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

segments. Thorax and tegulae grey-fuscous, some-
times grey irrorated with black. Forewing grey with
some bluish and purplish lustre; blackish brown post-
medial fascia distinctly oblique, not clearly demarcated,
but inwardly bounded by narrow zone of grey-white
scales; fascia with some gold lustre, similar scales
forming an irregular blackish brown spot at apex, just
before cilia. Cilia grey. Underside of forewing fuscous
brown or brown in apical half, but tending to be paler
in basal half. Hindwing and cilia grey or greyish. No
androconia on hindwing or forewing. Legs grey-brown
or fuscous brown on upperside, cream or ochreous
cream on underside. Abdomen fuscous on upperside,
ochreous brown or pale brown on underside; tufts
ochreous brown, hardly contrasting.

FEMALE. Unknown.

MALE GENITALIA (Figs 178, 179). Capsule 378 um
long. Tegumen very large, with two ventral lobes di-
rected inwardly. Pseuduncus short but narrow. Uncus
inverted V-shaped, with long anterior extensions.
Gnathos with long caudal process and well developed,
broad lateral arms; central plate small, anteriorly with
two close-set papillae. Valva (244 uum long) relatively
narrow, simple, without additional lobes or extensions.
The two inward-directed lobes in the centre of the
genital capsule are part of the tegumen, not the valvae.
Transtilla with broad transverse bar and huge parallel
processes. Juxta absent. Vinculum large, ventral plate
almost square anteriorly, without anterior emargination;
posterior margin weakly sclerotized, almost indiscern-
ible. Aedeagus 332 um, with 9 large spine-like sclerites:
7 well-sclerotized carinae and two separate spines
which probably represent cornuti.

BIOLOGY. Adults collected in April.

DIAGNOSIS. Among congeneric Neotropical species,
novenarius slightly resembles only laxibasis and
hispidus in external features. However, from the last it
clearly differs in the absence of androconia on the
hindwing and the significantly smaller wingspan; well-
preserved specimens can be distinguished from
laxibasis by the generally darker body coloration and
by the distinct fuscous apical mark in the forewing (in
laxibasis the forewing apex is irrorated with dark
scales but they do not form a distinct blackish spot).
The male genitalia of novenarius may be distinguished
amongst those of similar species by the very large and
truncate vinculum and by the unique configuration of
spines at the apex of the aedeagus. The form of the
vinculum and aedeagus in combination with the sim-
ple valva separate this from all other Acalyptris species.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The type series is
well-preserved, with the exception of one specimen,
which is stored in a gelatine capsule.

49

MATERIAL EXAMINED.
Holotype d, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 28970.

Paratypes, 5d, data as holotype, genitalia slide nos
29103, AD0301 6 (BMNH (3); VPU(2)).

REMARKS. The inner extensions of the vinculum are
not isolated from the main structure and do not form
the separate lateral apodemes so characteristic of many
Acalyptris; the lack of separation could be interpreted
as a plesiomorphy, but the extensions are in other
respects quite specialized, being narrowed inwardly.
They could be mistaken for processes of the valvae
which partially cover them in ventral view (a disadvan-
tage when a permanent preparation is examined).

46. Acalyptris lascuevella sp. n.
(Figs 50, 180, 181)

MALE. (Fig. 50). Forewing length: 2.0 mm. Wing-
span: 4.5 mm. Head: palpi cream; frontal tuft
orange-ochre; collar comprised of ochreous cream
piliform scales; eye-caps ochreous cream or yellowish
cream; antenna dark grey-brown, about 29-30 seg-
ments. Thorax, tegulae and forewing densely irrorated
with brown-black or fuscous scales; basal half of
forewing with dark scales smaller and less distinct,
scales in apical half very distinct and contrasting with
greyish cream background. Forewing with two cream
spots, one larger and costal, distinctly oblique, post-
medial, the other about one-half the size, tornal; dark
scales less dense in median area of forewing, giving a
patchy appearance, probably variable, and with some
asymmetry in the holotype. Cilia cream, greyish cream
at certain angles, with overlapping. fuscous scales at
apex, forming an irregular cilia-line. Underside of
forewing fuscous. Hindwing lanceolate, greyish; cilia
greyish. No androconia on hindwing or forewing. Legs
greyish cream with some irregular fuscous shading.
Upperside of abdomen blackish fuscous, genital seg-
ments pale ochre; underside brown, genital segments
ochreous.

FEMALE. Unknown.

GENITALIA 6 (Figs 180, 181). Capsule 230-235 um.
Dorsal plate of tegumen a moderately small sclerite
forming a bilobed pseuduncus with rounded, divergent
sparsely setose lobes. Uncus inverted V-shaped, with
caudal process. Gnathos with large pointed caudal
process, well developed lateral arms and central plate
rounded at anterior corners. Tegumen giving rise to a
pair of lateral rod-like processes along valvae. Valva
156-159 um long, straight, very narrow in apical
quarter, slightly bulged inward medially, and with
inward-directed basal process at about one-quarter.
Sublateral processes long, but transtilla absent. Juxta

50

(fused with posterior margin of vinculum?) a complex
and strongly sclerotized W-shaped plate with lateral
arms. Vinculum very short with broadly triangular
lateral lobes; anterior emargination shallow. Aedeagus
203-211 um long, very broad (ca. 83 um); no cornuti
visible on vesica.

BIOLOGY. Adults collected in April.

DIAGNOSIS. The white postmedian forewing mark-
ings differentiate /ascuevella from other Neotropical
Acalyptris; however this feature may not be reliable in
worn or aberrant specimens because irrorated wings
easily lose darker scales. The male genitalia are distin-
guished by an unusually specialized bilobed
pseuduncus, apically narrow valva and very broad
aedeagus without spine-like carinae.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL.
well-preserved.

The holotype is

MATERIAL EXAMINED.

Holotype 3, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 29119 (BMNH).

REMARKS. In general, the particular specializations
of the aedeagal carinae in Acalyptris are important
diagnostic features for most species. The neotropical
lascuevella is remarkable in that carinae are absent, a
presumed secondary loss that is parallelled in the
otherwise only distantly related Nearctic species A.
distaleus (Wilkinson).

47. Acalyptris bifidus sp. n.
(Figs 51, 182, 183)

MALE (Fig. 51). Forewing length: 1.9 mm. Wing-
span: 4.3 mm. Head: palpi cream; frontal tuft ochreous
orange, rather large; collar indistinct, comprised of
ochreous cream piliform scales; eye-caps yellowish
cream, large; antenna grey-brown, ca. 31—32 segments.
Thorax and tegulae cream densely irrorated with
grey-brown except for large anterior area of thorax
which is practically without brown scales. Forewing
ground-colour greyish cream densely irrorated with
dark-tipped scales which may be brownish, grey-brown
or fuscous, almost black; basal half of wing with dark-
tipped scales only slightly contrasting with ground
colour, darker scales more distinct in apical half, mostly
fuscous; dark scales forming relatively small postme-
dian and apical spots; dark scales also clearly visible
on cilia. Cilia cream. Underside of forewing brown.
Hindwing relatively narrow, lanceolate, grey; cilia
greyish. No androconia on hindwing or forewing.
Legs bright cream with blackish lateral shading. Col-
our of abdomen unknown.

R. PUPLESIS AND G.S. ROBINSON
Unknown.

GENITALIA 6 (Figs 182, 183). Genital capsule 340—
343 um long. Tegumen small with huge pseuduncus
bearing long and narrow lateral lobes, each shallowly
bilobed at apex. Uncus inverted V-shaped, with long
caudal process. Gnathos with large caudal process,
short and broad lateral lobes and large, rounded central
plate. Tegumen forming lateral rod-like processes be-
neath and roughly parallel to valvae. Valva 190-193 um
long, remarkably straight, tapered towards apex, distal
third very slender, without processes or lobes. Juxta
weakly sclerotized, with caudal extension and lateral
arms arising from broad base recessed into posterior
margin of vinculum. Vinculum with very short ventral
plate, long lateral lobes, and deep and semicircular
anterior emargination. Aedeagus 296-302 um, with
pair of narrow, strongly sclerotized, spine-like lateral
carinae. Vesica with numerous small and very small
cornuti.

FEMALE.

BIOLOGY. Adults collected in April.

DIAGNOSIS. The irrorated forewing gives bifidus a
superficial resemblance to other Neotropical Acalyptris
including fortis, martinheringi, species 29135,
unicornis, platygnathos, species 29140, tenuijuxtus,
bicornutus and aberrant specimens of other species.
The male genitalia are distinctive, however, in possess-
ing a pair of very sharp, straight and strong spine-like
distal carinae on the aedeagus, a large gnathos plate, a
two-pronged pseuduncus and very tapered valva. The
form of the pseuduncus, aedeagus and valva distin-
guish this from all other Acalyptris species.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype is
well-preserved, the wings not spread.

MATERIAL EXAMINED.

Holotype 3, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.1v.1998 (Puplesis & Hill),
genitalia slide no. 29133 (BMNH).

REMARKS. This is apparently a morphologically iso-
lated species characterized by a few distinct
autapomorphies, two of which — the pseuduncus proc-
esses and the distinctive carinae — prompted the
scientific name of this species.

48. Acalyptris trifidus sp. n.
(Figs 52, 184-186)

MALE (Fig.52). Forewing length: 2.2—2.4 mm. Wing-
span: 5.1-5.4 mm. Head: palpi cream; frontal tuft
very pale, yellowish cream; collar indistinct, comprised
of cream piliform scales; eye-caps cream; antenna
cream tinted ochre with irregular darker brown mark-
ings, ca. 29-30 segments. Thorax and tegulae cream.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

Forewing cream with dark ill-defined postmedian fas-
cia formed by brown and blackish to black scales;
tornal region of fascia always extended distally; a few
sparse brown or blackish scales medially and on cilia.
Cilia cream. Underside of forewing brownish cream.
Hindwing covered with cream-brownish androconial
scales; very long distinctively pale brown androconial
scales also overlap one-fifth length of cilia in basal two-
thirds of wing. Cilia cream. Legs cream with irregular
brownish shading. Abdomen entirely cream.

FEMALE.
hindwing.

GENITALIA 6 (Figs 184-186). Capsule 280 um long.
Tegumen small, with broad, broadly rounded
pseuduncus. Uncus inverted V-shaped, with three
caudally directed blunt processes. Gnathos with large
but relatively slender caudal process and broad caudally
bent lateral arms; central plate absent. Tegumen with
lateral rod-like sclerites underlying valvae along the
capsule. Valva 170-178 um long, slender and slightly
sinuous, tapered towards apex; without processes or
lobe. Transtilla with broad transverse bar and
well-developed, but not very long sublateral process.
Juxta absent, but posterior margin of vinculum form-
ing small caudally rounded plate between valval bases.
Ventral plate of vinculum very short but broad, with
triangular lateral lobes; anterior emargination shallow
but broad. Aedeagus 326-333 um long, very broad,
with two large, almost straight, horn-like lateral cari-
nae with sharp tips; other more indistinct features
include lobate apical carinae (including a long and
broad dorsal plate). Ventral carina single, protruding
some distance from tube of aedeagus with which it is
connected basally via a very broad joint.

Similar to male; no androconial scales on

GENITALIA §. [Association is tentative only.] Total
length about 617—646 pm. S8 and T8 broad, caudally
truncate. Anal papillae undeveloped. Apophyses
posteriores and anteriores very narrow, almost the
same length (ca. 138-146 um), apophyses anteriores
slightly curved inward and pointed anteriorly. Vestibu-
lum relatively broad, with large, posteriorly pointed,
Y-shaped sclerite. Corpus bursae ovoid with a large
oval signum on each side, each comprising large cen-
tral oval cells surrounded by comb-like pectination.
Accessory sac undeveloped; ductus spermathecae un-
known (missing in the single available preparation).

BIOLOGY. Adults collected in April.

DIAGNOSIS. A. trifidus may be distinguished among
congeneric Neotropical species by its oblique fuscous
postmedian forewing fascia and the long androconia
overlapping the hindwing cilia; it resembles only
dividua but the latter species has a darker forewing
fascia and a paler hindwing with no androconia. Its
external features in combination with the remarkable

51

three-pronged uncus serve to differentiate it from all
other Acalyptris species.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. While the holotype
is in good condition, two of three paratypes have
abraded forewings.

MATERIAL EXAMINED.

Holotype d, Belize: Chiquibul Forest Reserve, Las
Cuevas, 3—16.iv.1998 (Puplesis & Hill), genitalia slide
no. 29127 (BMNH).

Paratypes, 3d, data as holotype, genitalia slide nos
29138, ADO310 (BMNH (2); VPU (1)). Excluded
from paratype series: 392, genitalia slide no. 29142
(BMNH).

REMARKS. The three females (above) are excluded
from the paratype series as their association with the
males is tentative.

49. Acalyptris tenuijuxtus (Davis, 1978)
comb. n.

Microcalyptris tenuijuxtus Davis, 1978: 216, 217.

MALE. Described and figured by Davis (1978: 216,
ie se
FEMALE. Similar to male.

GENITALIA 6. Described and figured by Davis (1978:
216, figs 21-23).

GENITALIA °. Described and figured by Davis (1978:
216, fig. 33).

BIOLOGY. Adults collected in early October to late
November.
DIAGNOSIS. Among congeneric Neotropical species,

tenuijuxtus is externally similar to fortis, martinheringi,
species 29135, unicornis, platygnathos, species 29140,
bifidus, bicornutus and aberrant specimens of other
species in having an irrorated forewing. However, it
differs in the structure of the male genitalia from all of
these in the isolation of the apodemes from the vincu-
lum, the S5-lobed pseuduncus + uncus and the
anchor-shaped juxta (similar to that of unicornis but
with a longer caudal process). From the closely related
bifidus it differs in the absence of a pair of slender and
pointed carinae on the aedeagus and the narrower
juxta. The anchor-shaped juxta, the 5-lobed pseuduncus
+ uncus and the long lobes of the vinculum together
differentiate this from all other Acalyptris species.

DISTRIBUTION. USA (Florida Keys).

50. Acalyptris unicornis sp. n.
(Figs 53, 187-190)

MALE (Fig. 53). Forewing length: 1.8—1.9 mm. Wing-

52

span: 4.44.5 mm. Head: palpi cream; frontal tuft
orange-ochre; collar indistinct, comprised of pale
ochreous piliform scales; eye-caps cream; antenna
brownish, ca. 32 segments. Thorax predominantly
cream, with few greyish scales posteriorly; tegulae
cream. Forewing cream, irregularly irrorated with grey,
brownish grey and fuscous scales; one or two indistinct
dorsal cream spots (without dark scales). Cilia greyish
cream. Underside of forewing cream. Hindwing
upperside and underside cream, relatively narrow, lan-
ceolate; cilia concolorous. No androconia on hindwing
or forewing but the unusually pale colour of the
hindwing scales may indicate that they have an
androconial function. Legs cream with some greyish
shading at front. Abdomen cream on upperside and
underside.

FEMALE. Unknown.

GENITALIA 6 (Figs 187-190). Capsule 300-320 pm
long. Pseuduncus a narrow, weakly sclerotized, apically
papillate caudal extension of tegumen. Uncus very
complex, with ventrally directed long spine-like proc-
ess (Fig. 190 — ventral component) and two caudally
directed dorsal processes, each with a small lateral
papilla (Fig. 187 — not shown in Fig. 190). Gnathos
with large pointed caudal process, broad lateral arms
and weakly developed central plate that is almost
quadrate anteriorly. Tegumen with lateral rod-like
sclerites along valvae. Valva 158-182 pm long, slen-
der, gradually narrowed apically, without any lobes or
processes. Transverse bar of transtilla absent; sublateral
processes very slender and short. Juxta (formed by
caudal extension of vinculum or fused with vinculum)
triangular, pointed and sclerotized at apex and with
lateral arms. Vinculum with two large lateral lobes;
anterior emargination deep and rounded. Aedeagus
280 um long, with one very large cornutus which at
certain angles can appear to be a curved sclerotized
plate (Fig. 189), two large spine-like apical
sclerotizations and numerous tiny cornuti on the me-
dial and basal areas of the vesica.

BIOLOGY. Adult collected in April.

DIAGNOSIS. A. unicornis resembles externally sev-
eral congeneric Neotropical species with an irrorated
forewing: fortis, martinheringi, species 29135,
platygnathos, species 29140, bifidus, bicornutus,
tenuijuxtus and aberrant specimens of other species.
The male genitalia differ from those of all other species
in the combination of complex uncus (similar in out-
line to the pseuduncus of tenuijuxtus), specialized
gnathos with ventral process, caudally pointed undi-
vided juxta (the juxta is more elongate and apically
bifid in tenuijuxtus, and much more narrowly articu-
lated with the vinculum), evenly tapered valva (not
bulged at one-third as in tenuijuxtus) and the single

R. PUPLESIS AND G.S. ROBINSON

large cornutus in the apical third of aedeagus (appar-
ently absent in fenuijuxtus).

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype is in
good condition.

MATERIAL EXAMINED.

Holotype d, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 29134 (BMNH).

REMARKS. ‘The pseuduncus and uncus of unicornis,
as in a few other Acalyptris, are difficult to interpret as
the structures are adjacent and not easily distinguished
in a permanent preparation where they overly one
another. The simple, almost membranous pseuduncus
in unicornis is fairly typical for Acalyptris. The uncus
is frequently complex; the Neotropical bovicorneus,
trifidus and laxibasis, for example, have an elaborated,
strongly sclerotized uncus with caudally-directed proc-
esses similar to those of unicornis (Figs 166, 184, 191).
In unicornis, these processes each have a small lateral
papilla. The uncus in unicornis additionally has a
ventrally-directed spine (Fig. 190) which is otherwise
developed in the predominantly Asiatic shafirkanus-
group (see Puplesis, 1990). The single large cornutus is
not uniquely distinctive among world Acalyptris but is
nevertheless unusual and distinctive. Both features
represent autapomorphies for unicornis, and prompt
the scientific name of the species.

51. Acalyptris laxibasis sp. n.
(Figs 54, 191, 192)

MALE (Fig. 54). Forewing length: 2.0 mm. Wing-
span: 4.9 mm. Head: palpi cream; frontal tuft ochreous
orange; collar indistinct, cream, comprised of piliform
scales; eye-caps cream; antenna pale brown, almost
cream, ca. 30 segments. Thorax generally cream, pos-
terior region and tegulae irrorated with brown. Forewing
cream or pale yellowish cream irrorated with brownish
and grey brownish scales, which may look darker
depending upon angle of view; dark scales most densely
distributed at base of forewing, especially dark and
dense along costa; oblique blackish brown postmedian
fascia edged inwardly with grey-cream; apical fascia
pale yellowish cream with a group of brown scales
before cilia. Cilia cream to yellowish cream. Under-
side of forewing cream or pale brownish cream.
Hindwing and cilia greyish cream. No androconial
patches on hindwing or forewing. Legs pale greyish
cream with some brownish shading. Abdomen cream
on upperside and underside.

Unknown.

GENITALIA 6 (Figs 191-192). Capsule 325-328 um
long. Pseuduncus a narrow caudal process with four

FEMALE.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

apical setae. Uncus inverted V-shaped with two rela-
tively long subcaudally directed processes. Gnathos
with large caudal process, broad lateral arms and very
small rounded central plate. Tegumen with lateral
rod-like sclerites along valvae. Valva 218-220 um,
gradually narrowed towards apex, abruptly broadened
at base; no processes or lobes on valva. Transtilla with
long slender transverse bar and sublateral processes.
Vinculum short but with large triangular lateral lobes;
anterior emargination deep and rounded. Aedeagus
315-320 pm, broad, with three short lateral carinae.
Vesica with a single long rod-like sclerotization, a few
indistinct spine-like cornuti, and a compact triangular
group of needle-like cornuti surrounded by numerous
tiny cornuti.

BIOLOGY. Adult collected in April.

DIAGNOSIS. Resembles novenarius and hispidus in
the oblique dark forewing fascia but differs from the
latter species in the absence of hindwing androconia
and the considerably smaller wingspan; it differs from
novenarius in the generally paler body scaling and in
the apical mark being indistinct and fuscous whereas
in novenarius it is a distinct blackish spot. The male
genitalia of laxibasis differ from those of all other
species in the combination of abruptly broadened valval
base of valva, two-pronged uncus and very broad
aedeagus. This species is probably most closely re-
lated to unicornis, but may be easily distinguished by
the abruptly broadened valva, absence of large distinct
apical cornutus, and much simpler uncus.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The hindwings of
the holotype are detached and stored in a gelatine
capsule; head and forewing scaling is well-preserved.

MATERIAL EXAMINED.

Holotype 3, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 29128 (BMNH).

52. Acalyptris bicornutus (Davis, 1978)
comb. n.

Microcalyptris bicornutus Davis, 1978: 212-214.

MALE. Described and figured by Davis (1978: 212,
fig. 2).
FEMALE. Similar to male.

GENITALIA 6. Described and figured by Davis (1978:
212, figs 18-20).

GENITALIA 2. Described and figured by Davis (1978:
214, fig. 32).

BIOLOGY. Adults collected from late September to
late November.

35)

DIAGNOSIS. This species resembles externally sev-
eral other Neotropical congeners with an irrorated
forewing: fortis, martinheringi, species 29135,
platygnathos, species 29140, bifidus, unicornis,
tenuijuxtus and aberrant specimens of other species.
The male genitalia of bicornutus differ from all other
Acalyptris species in the extremely long and extremely
narrow lateral lobes of the vinculum.

DISTRIBUTION. USA (Florida Keys).

53. Acalyptris species 29135
(Figs 55, 193-195)

MALE (Fig. 55). Forewing length: 2.3 mm. Wing-
span: 5.2 mm. Head: palpi cream; frontal tuft rather
large, deep orange-ochre; collar comprised of cream
piliform scales; eye-caps cream with few brown scales;
antenna fuscous on upperside, ochreous cream on
underside, ca. 42 short segments; antenna short by dint
of reduced length of individual segments. Thorax,
tegulae and forewing irrorated with blackish scales
which are less distinctive on thorax and forewing base,
in apical half of forewing contrasting strongly with
greyish cream background; irregular spots on forewing,
especially in the middle and close to tornus or costa.
Cilia yellowish grey, distally almost white, cilia-line
formed by greyish scales. Underside of forewing fus-
cous. Hindwing lanceolate, creamy grey or brownish
grey (depending upon angle of view); cilia creamy
grey. No androconia on hindwing or forewing. Legs
greyish cream with irregular blackish lateral shading.
Abdomen greyish cream in upperside and underside.

FEMALE. Unknown.

GENITALIA 6 (Figs 193-195). Capsule 460-466 um
long. Dorsal plate of tegumen relatively small, but
caudally extended into a remarkably broad and trun-
cate pseuduncus. Uncus inverted V-shaped with caudal
process. Gnathos with large caudal process,
well-sclerotized lateral arms and small trapezoidal
central plate. Vinculum with lateral rod-like sclerites
more or less along valvae. Valva 244-252 um long,
very slender, just slightly broadened in basal half,
without lobes or processes. Transtilla with narrow and
relatively long sublateral processes and long trans-
verse bar. Juxta unknown (specimen damaged). Ventral
plate of vinculum short, 1.5x wider than long; lateral
lobes well-developed, more or less triangular, rounded
anteriorly; anterior emargination shallow. Aedeagus
291-299 um long, relatively narrow, with a few not
very distinctive carinae and a spine-like sclerite on
vesica.

BIOLOGY. Adult collected in April.

DIAGNOSIS. This species resembles externally other
Neotropical congeners with an irrorated forewing: for-

54

tis, martinheringi, bicornutus, platygnathos, species
29140, bifidus, unicornis, tenuijuxtus and aberrant
specimens of other species. However, the male genita-
lia of species 29135 differ from those of all Neotropical
species except martinheringi and bicornutus in the
caudally truncate pseuduncus; from martinheringi they
differ in the absence of a paired juxta and long and
sinuous spinose carinae on the aedeagus, and from
bicornutus in the short and rounded vinculum lobes.
The narrow, unarmed valva of this species is also
distinctive. The combination of the truncate pseuduncus
(parallelled in some other species) with the presence of
a transtilla and apodemes fused with the vinculum
separate this from all other Acalyptris.

DISTRIBUTION. Belize.

CONDITION OF MATERIAL. The single specimen is
well-preserved, but the genitalia are damaged and we
cannot be sure of the natural position of the valvae; the
juxta is missing and the aedeagus may not be intact.

MATERIAL EXAMINED.

Belize: 1d, Cayo District, Chiquibul Forest Reserve,
Las Cuevas, 3—16.iv.1998 (Puplesis & Hill), genitalia
slide no. 29135 (BMNH).

REMARKS. This distinctive species is described but is
not named because of the poor quality of the genitalia
preparation.

54. Acalyptris dividua sp. n.
(Figs 56, 196-198)

MALE (Fig.56). Forewing length: 2.2—2.3 mm. Wing-
span: 5.0—5.2 mm. Head: palpi cream to yellowish
cream; frontal tuft ochreous orange; collar cream,
indistinct; eye-caps yellowish or yellowish cream; ant-
enna yellow to pale ochreous yellow, ca. 28-30
segments. Thorax, tegulae and forewing yellowish or
yellowish cream; forewing with single subterminal
fascia oblique, broad, extended to cilia at tornus, formed
by densely distributed blackish scales. Cilia yellowish
cream. Underside of forewing cream. Hindwing rela-
tively broad, yellowish cream, cilia greyish cream to
greyish yellow. No androconia on forewing or
hindwing. Legs yellowish. Abdomen yellowish cream
to brownish on upperside, glossy cream or yellowish
on underside; anal tufts paired, piliform, relatively
long, cream; genital segments not contrasting in colour
with venter of abdomen.

FEMALE. Unknown.

GENITALIA 6 (Figs 196-198). Capsule 435-444 um
long. Pseuduncus broad, almost semicircular, papil-
lated. Uncus with two long divergent caudal processes,
each with a few setae. Tegumen simple, relatively
large. Gnathos with shallow U-shaped transverse bar
on very long and slender lateral arms. Valva ca. 265 um

R. PUPLESIS AND G.S. ROBINSON

long, with pointed, inward-directed median spine-like
process and slender apical process; base of valva
abruptly broadened. Transtilla with transverse bar and
well-developed but not very long sublateral processes.
Vinculum long, anteriorly very broadly rounded, with-
out lateral lobes. Aedeagus ca. 341 pm long, with three
very large horn-like cornuti extending beyond apex
(Figs 197, 198). Juxta indistinct, a semicircular mem-
branous lobe.

BIOLOGY. Adults collected in April.

DIAGNOSIS. This is an externally very distinctive
species in the Neotropical context, resembling only
trifidus with an oblique fuscous forewing postmedian
fascia. In dividua the forewing fascia is darker and
there are no long androconia overlapping the hindwing
cilia. The U-shaped gnathos, divided uncus and long
apical sclerites of the aedeagus are a unique combina-
tion, distinguishing this from all other Acalyptris.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The type series is
well-preserved, but one paratype is slightly abraded;
the wings of both specimens are not spread so the
hindwings are obscured.

MATERIAL EXAMINED.
Holotype d, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 29125 (BMNH).

Paratypes, 2, data as holotype, genitalia slide no.
AD039 (BMNH; VPU).

REMARKS. The species exhibits numerous distinc-
tive features, in most cases autapomorphic. The
scientific name refers to the divided uncus. The shape
of the gnathos is quite atypical for Acalyptris and sets
dividua apart from its congeners.

55. Acalyptris platygnathos sp. n.
(Figs 57, 199-202)

MALE (Fig. 57). Forewing length: about 1.7 mm.
Wingspan: 3.9-4.0 mm. Head: palpi greyish cream:
frontal tuft greyish ochre; collar greyish cream, com-
prised of piliform scales; eye-caps ochreous cream;
antenna grey-brown (creamy brown at certain angles),
ca. 29-30 segments. Thorax, tegulae and forewing
with greyish cream background densely irrorated with
greyish to fuscous brown or fuscous scales which are
most distinctive at apex and at middle of forewing
closer to dorsal margin; forewing dorsum with elon-
gate cream area one-third wing length without grey or
fuscous scales; costal margin with irregular pale me-
dian area. Cilia and underside of forewing grey-brown.
Hindwing lanceolate but not very slender; grey; cilia

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

grey. No androconia visible on hindwing or forewing.
Legs greyish or creamy grey with some irregular
fuscous markings. Abdomen grey or brownish cream.

FEMALE. Unknown.

GENITALIA 6 (Figs 199-202). Capsule 222-230 um
long. Tegumen moderately large, extended caudally
into a broad and very shallowly bilobed pseuduncus
(Fig. 199). Uncus reduced, fused with pseuduncus
(Fig. 199). Gnathos with an extraordinary broad, quad-
rate, plate-like caudal process with deep posterior
emargination; lateral arms of gnathos well-developed
and sclerotized; central plate, as such, absent (Fig. 201).
Valva 180-185 um long, more or less triangular in
ventral view, with narrow, slightly incurved apex and
papillated inner margin; with distinct bidentate medial
process and huge, caudally curved horn-like process
from dorsal surface at one-third. Transtilla with nar-
row transverse bar and slender, moderately long
sublateral processes; valvae joined at their bases by
membranous ventral connection. Vinculum large, ven-
tral plate trapezoidal with broadly rounded corners,
without lobes or emargination. Aedeagus 265-270 um
long, with eight almost symmetrical carinae; dorsal
pair connected by strong transverse bar-like
sclerotization, shorter lateral carinae appearing fused
with dorsal carinae because they overlap; ventral
spine-like carinae each with a smaller spinose carina at
base (Fig. 200). Vesica with very many tiny cornuti.

BIOLOGY. Adults collected in April.

DIAGNOSIS. This species resembles externally other
Neotropical congeners with an irrorated forewing: for-
tis, martinheringi, bicornutus, species 29135, species
29140, bifidus, unicornis, tenuijuxtus and aberrant
specimens of other species. The male genitalia, how-
ever, are distinctive, exhibiting a uniquely broad
gnathos and characteristic broad vinculum. The only
other Acalyptris species with a broad gnathos is the
southern European maritima Lastuvka & Lastuvka,
with a triangular rounded gnathos, unarmed valva and
tiny vinculum.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype has a
slightly abraded frontal tuft and left forewing.

MATERIAL EXAMINED.

Holotype d, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 29132 (BMNH).

REMARKS. Extremely ordinary externally, this small
species possesses comparatively large male genitalia
that are morphologically extreme, the valva with two
internal processes, an unusually broad vinculum, a
bench-shaped pseuduncus and unique, broad, slab-
shaped gnathos. These autapomorphies are parallelled

55

by another feature of interest in the context of Acalyptris
morphology as a whole — the uncus is reduced, and
partly fused with the pseuduncus (Fig. 199).

56. Acalyptris species 29140
(Figs 4-6, 58, 221)

MALE. Forewing length: 1.7—1.8 mm. Wingspan:
3.9-4.1 mm. Head: palpi cream; frontal tuft very pale,
ochreous orange; collar indistinct, cream; eye-caps
cream; antenna brownish cream, ca. 28 segments. Tho-
rax and tegulae yellowish cream, slightly glossy.
Forewing same colour as thorax but irrorated distally
with brownish and further distally with black. Cilia
cream. Distal area of forewing underside covered with
blackish androconial scales, some blackish scales also
along costal margin, elsewhere cream. Hindwing
cream, lanceolate, slender; cilia cream. No androconia
on hindwing. Legs cream with blackish shading. Ab-
domen brownish cream.

FEMALE (Fig. 58). Antenna ca. 23-24 segments.
Distal area of forewing underside without blackish
androconial scales, underside uniformly cream. Other-
wise as in male.

GENITALIA 6. Unknown.

GENITALIA 2(Fig. 221). Total length about 676 um.
S8 and T8 broad, caudally truncate. Anal papillae
undeveloped. Apophyses posteriores and anteriores
almost same length (ca. 130 um), but apophyses
anteriores two to three times thicker. Vestibulum rela-
tively broad, with large Y-shaped sclerite. Corpus
bursae oval with an oval signum on each side. Acces-
sory sac undeveloped; ductus spermathecae very
slender and short, sinuous and sclerotized.

BIOLOGY. Hostplant: Lonchocarpus lineatus Pittier
(Leguminosae — tree). Mines in leaves (Figs 4-6). Egg
grey-brown, laid on upperside of leaf close to a rib.
The larval mine starts as a long, very gradually broad-
ening sinuous or contorted gallery, initially filled with
dark brown frass, thereafter frass loosely dispersed in
a broad band leaving narrow clear margins. Larva pale
yellowish with brown central line. Cocoon ochreous
brown, 1.6—1.8 mm long, 1.0—1.2 wide. Adults col-
lected in April. In Belize isolated larvae occur along a
wide forest track.

DIAGNOSIS. This species resembles to some extent
other Neotropical congeners with an irrorated forewing:
fortis, bicornutus, platygnathos, species 29135, bifidus,
unicornis, tenuijuxtus and aberrant specimens of other
species. But it most closely resembles another pale
Neotropical species, Acalyptris martinheringi. From
all of these it tends to differ in the distinct predomi-
nance of dark scales in the apical half of the forewing
while the base remains almost entirely pale; in

56

martinheringi dark scales are distributed everywhere.
Diagnosis is complicated because the male genitalia
are unknown; key characters for identification are
currently no more than the apical distribution of dark
scales of forewing and Lonchocarpus lineatus Pittier
(Leguminosae) as a hostplant.

DISTRIBUTION. Belize.

CONDITION OF MATERIAL AVAILABLE. The single
available male lacks an abdomen; the remaining female
specimens are in poor condition, one in a gelatine
capsule, another with a mouldy head, and the third
with a single loose hindwing.

MATERIAL EXAMINED.

Belize: 1d, 32, Cayo District, Chiquibul Forest Re-
serve, Las Cuevas, 3—16.iv.1998, larvae on
Lonchocarpus lineatus Pittier, ex. 1. 26—30.iv.1998
(Puplesis & Hill), forewing venation slide 291444,
genitalia slides nos 291409, AD3122 (BMNH (14,
2 2); VPU (1 9); leaf-mine herbarium sheet 4431 (VPU).

GLAUCOLEPIS Braun

57. Glaucolepis aerifica (Meyrick, 1915)
comb. n.

Nepticula aerifica Meyrick, 1915: 255.
Stigmella aerifica (Meyrick), Davis, 1984: 18.

(Figs 59, 203, 204, 222)

MALE (Fig. 59). Forewing length: 2.7 mm. Wing-
span: 5.8 mm. Head: palpi greyish; frontal tuft greyish
yellow, very pale; collar indistinct; eye-caps greyish
white, relatively small; antenna brown, 36-38 seg-
ments. Thorax and tegulae shining greenish bronze.
Forewing smoothly scaled before fascia, shining green-
ish bronze, with gold lustre at certain angles, brown
beyond fascia purple lustre; fascia poorly defined,
distinctly postmedian, almost terminal, shining sil-
very. Cilia brownish. Underside of forewing brown.
Hindwing relatively long, brown; cilia pale brown. No
androconia visible on forewing or hindwing. Colour
of abdomen unknown, most likely brown.

FEMALE. Forewing before fascia dark grey brown
with gold and light purple lustre. Scaling coarser than
in male. Otherwise similar to male, assuming the
female is conspecific (see Remarks).

GENITALIA 6 (Figs 203, 204). Capsule 410-415 um
long. Uncus band-shaped (slightly unnaturally turned
ventrad in Fig. 203, but this is not the natural position),
and extended caudally, papillate laterally. Tegumen
very short, simple. Gnathos with very broad trapezoidal
caudal process and long lateral arms. Valva 220 um
long, with long pointed apical process, with curved,

R. PUPLESIS AND G.S. ROBINSON

pointed inward-directed process at one-third. Trans-
verse bar of transtilla interrupted medially, sublateral
process triangular, tapered anteriorly. Juxta complex,
with long pointed caudal lobes and transversely
sclerotized base. Vinculum very large, anteriorly
rounded and slightly but gradually narrowed, without
emargination or lateral lobes. Aedeagus 290-295 um
long, with two rod-like sclerites fused caudally and
two membranous caudal processes with numerous
very fine spines; these processes are connected to the
aedeagus wall and are not part of the vesica. Vesica
with a few pointed and rounded sclerotizations.

GENITALIA @ (Fig. 222). Total length 976-980 um.
T8 triangular. Anal papillae not developed. Apophyses
anteriores broad and blunt anteriorly. Apophyses
posteriores long (260-265 pm) and slender, as long as
apophyses posteriores. Vestibulum relatively narrow,
without pectinations or sclerites. Caudal part of corpus
bursae coarsely folded, anterior region broadly oval,
densely covered with pectinations, without signa, but
pectinations at anterior end of corpus bursae tending to
form one or two continuous transverse lines. Acces-
sory sac coarsely folded, broad; ductus spermathecae
slender, membranous except for a ring-shaped sclerite
some distance from accessory sac.

BIOLOGY. Adults collected in July.

DIAGNOSIS. This species differs from other known
representatives of Glaucolepis and indeed all
nepticulids either by the combination of the distinctly
postmedial forewing fascia, the broad and bilobed
juxta, broad gnathos and unique morphology of the
aedeagus. This is an outstanding species, whose taxo-
nomic position is ambiguous and which will repay
more detailed study when further material becomes
available.

DISTRIBUTION. Peru.

CONDITION OF TYPE MATERIAL. The lectotype is badly
damaged: the head is on a pin and shows a rubbed
frontal tuft and indistinct collar; the thorax and wings
are pinned separately in a gelatine capsule; the scaling
of thorax and wings is well preserved; damage to the
thorax precludes repinning. The paralectotype is
slightly rubbed but more or less satisfactorily pinned.

MATERIAL EXAMINED.

Lectotype d, Peru: Oroya, vii.1914 (Parish), genitalia

slide no. 28965 6 (BMNH), here designated.
Paralectotype 2, data as lectotype, genitalia slide no.

28966 2 (BMNH).

REMARKS. This species’ taxonomic position is un-
certain and provisional. The male genitalia possess a
transverse transtilla bar which is generally uncharac-
teristic of Glaucolepis. However, the transtilla is
interrupted in the middle. Some Glaucolepis, such as

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

the raikhonae-group (Nieukerken & Puplesis, 1991),
have partially or completely reduced signa. On the
another hand, the shape of sclerites in the aedeagus
resemble those of Glaucolepis. The wing venation of
this species has not been studied in order to preserve
the already badly damaged lectotype.

The darker forewing colour and coarser scaling of
the female paralectotype in comparison with the male
holotype gives us pause in assuming that the two
represent opposite sexes of the same species. More-
over, the female genitalia have no signa, atypical of
Glaucolepis. If the female is not aerifica, then it might
not even be a Glaucolepis. However, the distinctly
postmedial fascia of the forewing (a character gener-
ally uncommon amongst nepticulids) is shared by both
specimens.

58. Glaucolepis argentosa sp. n.
(Figs 60, 205, 206)

MALE (Fig. 60). Forewing length: 2.0—2.2 mm. Wing-
span: 4.5-4.7 mm. Head: palpi cream; frontal tuft
orange-ochre; collar comprised of ochreous cream to
cream piliform scales; eye-caps ochreous cream to
orange-cream; antenna brown, ca. 32 segments. Tho-
rax and tegulae with dark metallic or silvery lustre.
Forewing dark blackish brown, practically black
apically, very glossy, with some gold lustre, basal third
of forewing and very distinct postmedial fascia silver,
glossy; silvery fasciae connected along dorsal margin
of forewing. Cilia dark grey, pale-tipped, glossy silver,
usually with distinct cilia line of black scales. Under-
side of forewing grey to fuscous. Hindwing narrow,
lanceolate, greyish cream; cilia cream or greyish cream.
No androconial patches on hindwing or forewing.
Legs ochreous cream with brown to blackish lateral
shading. Abdomen upperside blackish, no tufts visible,
genital segments ochreous or yellowish cream, indis-
tinct; underside orange-cream or ochreous cream with
distinct concolorous genital segments (valvae).

FEMALE. Unknown.

GENITALIA 6 (Figs 205, 206). Capsule 346-353 um
long. Tegumen very large, caudally extended into a
short and narrow pseuduncus which is fused with
uncus. Uncus represented by two lateral processes,
each with one large ventral seta. Gnathos with large
caudal process and broad, strongly sclerotized, distally
elaborated lateral arms; central plate weakly devel-
oped, with shallow anterior emargination. Valva
230-246 um long, very broad, quadrate in basal half,
tapered towards lateral margin in apical half; apical
process hardly developed, represented by a small den-
tate process; apical margin with three long stout setae
and numerous slender setae. Valvae joined basally by
strong ventral connection that functionally replaces
the transverse transtilla bar. Transtilla absent, i.e., no

o7

transverse bar; sublateral processes (i.e., valval
apodemes) straight, elongate. Vinculum very large and
broad, ventral plate slightly tapered anteriorly, without
anterior emargination or lateral lobes. Aedeagus 311—
323 um, very broad (approximately 146 um in middle),
with two sclerotized, distally pointed apical carinae
and a short rounded lobe; with two very long and
narrow longitudinal rod-like sclerotizations parallel to
tube walls; vesica with many moderately or weakly
sclerotized, spine-or needle-like cornuti concentrated
towards apex of aedeagus; single very large, horn-like
cornutus centrally, weakly sclerotized and indistinct;
sparse sclerotization forming ill-defined cornuti vis-
ible in central part of vesica.

BIOLOGY. Adults collected in April.

DIAGNOSIS. The combination of unusual forewing
pattern with glossy silver markings, together with the
paired elongate sclerites in the aedeagus and the bifid
uncus make this species unique among all nepticulids.

DISTRIBUTION. Belize.

CONDITION OF TYPE MATERIAL. The holotype is
well-preserved, with distinct wing pattern; most of the
paratypes have at least slightly abraded forewings; the
abdomen of one paratype is missing.

MATERIAL EXAMINED.
Holotype d, Belize: Cayo District, Chiquibul Forest
Reserve, Las Cuevas, 3—16.iv.1998 (Puplesis & Hill),
genitalia slide no. 29106d (BMNH).

Paratypes, 6d, data as holotype, genitalia slide no.
AD0303 (BMNH (4); VPU (2)).

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INDEX

Synonyms and misidentifications are in italics; principal references are in bold.

ACALYPTRIS 16, 18, 45
aerifica 18, 56
albilamina 10, 17, 33, 34
andina 10, 17, 24
argentosa 18, 57 gossypii 13, 17, 35
guittonae 18, 40
barbata 11, 18, 37

basidactyla 4, 7, 17, 22 hamata 17, 30
bicornutus 18, 53 helenella 18, 41
bifidus 18, 50 hilli 17,19

hispidus 18, 48
hylomaga 18, 39

bovicorneus 11, 18, 45

castaneaefolia 18
clemensella 18
contracoloreal7, 20
costalimai 18, 40
cuprata 17, 24
cyanochlora 4, 10, 17, 18

imperatoria 17, 30
johannis 4, 17, 25
kimae 17, 35

lascuevella 18, 49
latipennata 18, 45
laxibasis 18, 52

diskusi 18, 43
dividua 18, 54

ECTOEDEMIA 16, 18, 41
ENTEUCHA 15, 17, 18
epicosma 7, 17, 28
eurydesma 17, 32
eurydesma group 11, 17, 32

marmorea 17, 26
mesoloba 18, 41

myricafoliella 18
FOMORIA 16, 18, 43

fortis 18, 47
fuscilamina 10, 17, 33, 34

nigriverticella 18
novenarius 18, 48

fuscivittata 7, 11, 18, 42

gilvafascia 13, 17, 19
GLAUCOLEPIS 17, 18, 56

martinheringi 18, 46

molybditis 4, 18, 43

obrutella 18
olyritis 17, 31
ostryaefoliella 18
ovata 7, 10, 18, 39

peruanica 17, 27
platygnathos 11, 18,54
plumosetaeella 4, 17, 36
pruinosa 10, 18, 38

reneella 18, 41
rudis 17, 26

salicis group 11, 17
schoorli 17, 29

similella 18

snaddoni 17, 21

species 29105 10, 13, 18, 41
species 29122 11, 18, 44
species 29135 18, 53
species 29140 13, 18, 55
STIGMELLA 16, 17, 23

MANONEURA 15, 17, 22

tenuijuxtus 18,51
terricula 17, 20
trifidus 10, 18, 50
trinaria 17, 23

unicornis 18, 51

virgulae 18

60 R. PUPLESIS AND G.S. ROBINSON

Figs 4-6 Mines of Acalyptris species 29140 on Lonchocarpus lineatus Pittier (Leguminosae). 4, twig with two sinuous
mines on one leaf; 5, leaf with two contorted mines; 6, gallery showing frass distribution.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

Figs 7-10 Enteucha species. 7, cyanochlora 3 (Guyana); 8, hilli 3 (Belize); 9, contracolorea 3 (Belize); 10, terricula 3
(Peru).

61

62 R. PUPLESIS AND G.S. ROBINSON

11

12

13

a pif 4 -
‘elt | ‘ | Wi oon

Se
—
=
ee
——-

14

Figs 11-14 Neotropical Nepticulidae. 11, Enteucha snaddoni ? (Belize); 12, Manoneura basidactyla 3 (Belize); 13,
Manoneura trinaria 3 (Venezuela); 14, Stigmella andina ¢ left side, ? right side (Peru).

63

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

Figs 15-18 Stigmella species. 15, cuprata 3 (Peru); 16, johannis 3 (Colombia); 17, rudis 3 (Argentina); 18, marmorea

6 (Peru).

R. PUPLESIS AND G.S. ROBINSON

64

Figs 19-24 Stigmella species (Peru). 19, peruanica 3; 20, epicosma, lectotype d (Lima, 500 ft); 21, epicosma 9, not

type series (Rio Andamayo Valley, ca. 3000 m); 22, hamata 3; 23, imperatoria 3; 24, schoorli 6d.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

|

uh

Bye
E==

fh

Figs 25-30 Stigmella species. 25, eurydesma 3 (Guyana); 26, albilamina 3 (Belize); 27, fuscilamina 3 (Belize); 28,
olyritis 3 (Peru); 29, gossypii 3 (after Newton & Wilkinson, 1982) (Puerto Rico); 30, kimae, 3 (Belize).

65

R. PUPLESIS AND G.S. ROBINSON

66

Figs 31-35 Stigmella species. 31, plumosetaeella 3 (Mexico); 32, barbata d (Belize); 33, pruinosa @ left side, 3 right side

(Belize), 34, pruinosa, male hindwing (Belize); 35, ovata d (Argentina).

67

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

|

I

Neotropical Nepticulidae. 36, Stigmella hylomaga 2? (Argentina); 37, S. costalimai 2 (Argentina); 38,

S. guittonae 2 (Argentina); 39, Ectoedemia fuscivittata 3, (Belize); 40, Ectoedemia species 29105 3 (Belize),

Figs 36-40

68 R. PUPLESIS AND G.S. ROBINSON

41

42

43

44

Figs 41-44 Fomoria species. 41, molybditis 5 (Colombia); 42, diskusi 3 (Belize); 43, latipennata 3 (Belize); 44, species
29122 6, reconstructed (Belize).

69

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

4

46
47
48

it! HH

WE
) Wy, Wy. (a,
—— ———— Ree ==

———— ===
See = : Fs Ti
— ===
——————— —
——— i —— Re SS
SS 4 = Ry ——S
———=S> f ee — yet ———_—
——> OM 3 Fy ———— Hasee i] — —
SS>= i. ——— een ///, —
— Mey / ; a ==
ve , WSS

: My —— tt
. \ / i. ZEN iil
aN |

Figs 45-48 Acalyptris species, males (Belize). 45, bovicorneus; 46, martinheringi; 47, fortis; 48, hispidus.

R. PUPLESIS AND G.S. ROBINSON

70

52

Figs 49-52 Acalyptris species, males (Belize). 49, novenarius; 50, lascuevella; 51, bifidus; 52, trifidus.

i

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

Yy

My

Rin noes

|
4

lL
Ww
5

\

Poa

oe Oe

f A ¢ Yi
| —— eels ey, Y
LL, =S= / 2

Figs 53-56 Acalyptris species, dd (Belize). 53, unicornis; 54, laxibasis; 55, species 29135; 56, dividua.

R. PUPLESIS AND G.S. ROBINSON

72

57

Wp,

SSS—_
S———

————————

Pak

Figs 57-60 Neotropical Nepticulidae. 57, Acalyptris platygnathos 3 (Belize); 58, Acalyptris species 29140 Q, left &

right sides (Belize); 59, Glaucolepis aerifica 3 (Peru); 60, Glaucolepis argentosa 3 (Belize).

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 73

Sc Ry +243

A 65

Figs 61-65 Wing venation. 61, Manoneura basidactyla, 3 forewing & hindwing; 62, Stigmella ovata, d forewing; 63,
Stigmella albilamina, 2 forewing and hindwing; 64, Fomoria diskusi, 3 forewing; 65, Acalyptris bovicorneus, 3
forewing. Scale: 0.5 mm.

74 R. PUPLESIS AND G.S. ROBINSON

ventral
lobe

Figs 66-70 Male genitalia of Enteucha species. 66, cyanochlora, holotype (Guyana) (12273 — BMNH), gnathos; 67, same,
aedeagus; 68, same, capsule; 69, hilli, holotype, Belize (28967 - BMNH), capsule; 70, same, aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 12

tegumen

sublateral
process

Figs 71-74 Male genitalia of Enteucha contracolorea, holotype, Belize (29114 —- BMNH). 71, capsule; 72, uncus and
tegumen; 73, gnathos; 74, aedeagus. Scale: 0.1 mm.

76 R. PUPLESIS AND G.S. ROBINSON

gnathos

sublateral
process

75

Figs 75-79 Male genitalia of Enteucha terricula, holotype, Peru (Diskus183—-ZMUC). 75, capsule; 76, aedeagus; 77, uncus
and tegumen; 78, gnathos; 79, valva (apical process omitted). Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE V7

unCUS

tegumen

gnathos

82

Figs 80-82 Male genitalia of Enteucha snaddoni, holotype, Belize (29117 — BMNH). 80, capsule; 81, uncus and tegumen;
82, aedeagus. Scale: 0.1 mm.

78 R. PUPLESIS AND G.S. ROBINSON

uncus

: tegumen
apical

process

tegumen

gnathos

84

+ carina

carina =

sublateral
process

85

Figs 83-85 Male genitalia of Manoneura basidactyla, Belize (29120 — BMNH). 83, capsule; 84, gnathos, uncus and
tegumen; 85, aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 1S)

tegumen

ventral
carinae

IP] f
sublateral i;
process ee

Figs 86, 87 Male genitalia of Manoneura trinaria, holotype, Venezuela (Diskus003 — USNM). 86, capsule; 87, aedeagus.
Scale: 0.1 mm.

80 R. PUPLESIS AND G.S. ROBINSON

Figs 88-91 Male genitalia of Stigmella species. 88, andina, lectotype, Peru (20612 - BMNH), capsule; 89, same, aedeagus; 90,
cuprata, lectotype, Peru (28848 — BMNH), capsule without valvae; 91, same, valvae and aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

Figs 92-96 Male genitalia of Stigmella species. 92, johannis, holotype, Colombia (28843 — BMNH), capsule; 93, same,
aedeagus; 94, rudis, holotype, Argentina (Diskus175 — ZMUC), aedeagus; 95, same, paratype (DiSkus180 — ZMUC),
characteristic cluster of cornuti on shared base; 96, same, holotype (Diskus175 — ZMUC), capsule. Scale: 0.1 mm.

81

82 R. PUPLESIS AND G.S. ROBINSON

uncus

Figs 97,98 Male genitalia of Stigmella marmorea, holotype, Peru (Diskus182 —ZMUC). 97, capsule; 98, aedeagus. Scale:
0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 83

Figs 99-103 Male genitalia of Stigmella epicosma, Peru. 99, lectotype: Lima (28849 — BMNH), valvae, unnaturally spread;
100, same, capsule; 101, same, aedeagus; 102, non-type male, Rio Andamayo Valley (Diskus187 — ZMUC), aedeagus; 103,
same, capsule. Scale: 0.1 mm.

84 R. PUPLESIS AND G.S. ROBINSON

Figs 104-107 Male genitalia of Stigmella species, Peru. 104, peruanica, holotype (Diskus189 — ZMUC), capsule; 105,
same, aedeagus; 106, schoorli, holotype (Diskus200 — ZMUC), capsule; 107, same, aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

Figs 108-112 Male genitalia of Stigmella hamata, holotype, Peru (Di8kus199 — ZMUC). 108, capsule; 109, uncus and
tegumen; 110, left valva; 111, aedeagus; 112, gnathos. Scale: 0.1 mm.

85

86 R. PUPLESIS AND G.S. ROBINSON

Figs 113-116 Male genitalia of Stigmella imperatoria, holotype, Peru (Diskus195 — ZMUC). 113, capsule; 114, uncus
and tegumen; 115, gnathos; 116, aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

cornutus

apical
process

Figs 117-122 Male genitalia of Stigmella albilamina, Belize. 117, holotype (29113 - BMNH), capsule; 118, same,
aedeagus; 119, paratype (29113 - BMNH), left valva; 120, same, gnathos and uncus; 121, same, juxta; 122, same,
aedeagus. Scale: 0.1 mm.

87

88 R. PUPLESIS AND G.S. ROBINSON

124

apical
process

uncus

127

Figs 123-127 Male genitalia of Stigmella fuscilamina, Belize. 123, holotype (29110 —BMNH), capsule; 124, same,
aedeagus; 125, paratype (29111 — BMNH), left valva; 126, same, gnathos and uncus; 127, same, aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

apical
process

129

apical 130

process

132

7) 131

Figs 128-132 Male genitalia of Stigmella species. 128, eurydesma, lectotype, Guyana (28842 — BMNH), capsule; 129,

same, gnathos and uncus; 130, same, aedeagus; 131, olyritis, lectotype, Peru (28851 — BMNH), capsule; 132, same,

aedeagus. Scale: 0.1 mm.

89

90 R. PUPLESIS AND G.S. ROBINSON

apical
process

apical
process

Figs 133-138 Male genitalia of Stigmella species. 133, gossypii, Puerto Rico (after Newton & Wilkinson, 1982), capsule;
134, same, aedeagus; 135, kimae, holotype, Belize (29118 - BMNH), valva; 136, same, transtilla; 137, same, capsule (with
replaced and slightly squashed valvae); 138, same, aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

143

Figs 139-143 Male genitalia of Stigmella barbata, holotype, Belize (29137 — BMNH). 139, gnathos; 140, valva and
enlarged seta; 141, capsule; 142, aedeagus; 143, uncus and tegumen. Scale: 0.1 mm.

91

92 R. PUPLESIS AND G.S. ROBINSON

Figs 144-147 Male genitalia of Stigmella species. 144, plumosetaeella, Mexico (Diskus005 — USNM), capsule; 145, same,
aedeagus; 146, pruinosa, holotype, Belize (29124 - BMNH), capsule; 147, same, aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

uncus

tegumen 4

transtilla
bar

Figs 148-151 Male genitalia of Stigmella ovata, holotype, Argentina (Di8kus184 —ZMUC). 148, gnathos, uncus and

tegumen; 149, juxta; 150, capsule; 151, aedeagus. Scale: 0.1 mm.

93

94 R. PUPLESIS AND G.S. ROBINSON

pseuduncus

carinae

cornuti

152

pseuduncus

Figs 152-156 Male genitalia of Ectoedemia species, Belize. 152, species 29105 (29105 — BMNH), capsule; 153, same,
aedeagus; 154, fuscivittata, holotype (29107 — BMNH), capsule; 155, same, aedeagus; 156, fuscivittata, paratype
(AD0302 — VPU), aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 95

uncus

ventral
process

transtilla
bar

Figs 157,158 Male genitalia of Fomoria molybditis, holotype, Colombia (25651 — BMNH). 157, capsule; 158, aedeagus.
Scale: 0.1 mm.

96

pseuduncus

uncus

Fi /]\ ventral
H\ LT processes

transtilla
bar

159

R. PUPLESIS AND G.S. ROBINSON

| cathrema

160

Figs 159, 160 Male genitalia of Fomoria diskusi, holotype, Belize (28844 — BMNH). 159, capsule; 160, aedeagus. Scale:

0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 97

uncus

ventral
lobe

cornutus

161

transtilla
bar

ventral
lobe

163

Figs 161-163 Male genitalia of Fomoria species 29122, Belize (29122 - BMNH). 161, capsule; 162, aedeagus with
partially everted vesica; 163, valva. Scale: 0.1 mm.

98 R. PUPLESIS AND G.S. ROBINSON

pseuduncus

Figs 164,165 Male genitalia of Fomoria latipennata, holotype, Belize (28968 — BMNH). 164, capsule; 165, aedeagus.
Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 99

pseuduncus

Figs 166-169 Male genitalia of Acalyptris bovicorneus, Belize. 166, holotype (29115 — BMNH), capsule; 167, same,
aedeagus; 168, paratype (AD0307 — VPU), aedeagus; 169, holotype (29115 — BMNH), aedeagus in lateral view. Scale:
0.1 mm.

100 R. PUPLESIS AND G.S. ROBINSON

Figs 170-173 Male genitalia of Acalyptris species, Belize. 170, martinheringi, holotype (29109 — BMNH), capsule; 171,
same, aedeagus; 172, hispidus, holotype (29104 — BMNH), capsule; 173, same, aedeagus. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 101

uncus pseuduncus

carinae

lateral
apodeme

174

sublateral
process

Figs 174-177 Male genitalia of Acalyptris fortis, Belize. 174, holotype (29131 — BMNH), capsule; 175, same, aedeagus;
176, paratype (AD0311 — VPU), valva, slightly squashed; 177, paratype (29136 — BMNH), uncus. Scale: 0.1 mm.

102 R. PUPLESIS AND G.S. ROBINSON

pseuduncus

lateral
apodeme
179
4

178

Figs 178,179 Male genitalia of Acalyptris novenarius, holotype, Belize (28970 — BMNH). 178, capsule; 179, aedeagus.
Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 103

qr pseuduncus

Figs 180-183 Male genitalia of Acalyptris species, Belize. 180, lascuevella, holotype (29119 - BMNH), capsule; 181,
same, aedeagus; 182, bifidus, holotype (29133 - BMNH), capsule; 183, same, aedeagus. Scale: 0.1 mm.

104 R. PUPLESIS AND G.S. ROBINSON

ventral
process

Figs 184-190 Male genitalia of Acalyptris species, Belize. 184, trifidus, holotype (29127 BMNH), capsule; 185, same,
aedeagus; 186, same, ventral projection of aedeagus in lateral view; 187, unicornis, holotype (29134 — BMNH), capsule;
188, same, aedeagus; 189, cornutus in lateral view — compare 188; 190; same, ventral component of uncus, ventrolateral
view (dorsal component not shown — see 187). Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 105

uncus

pseuduncus

ae

sublateral
process

192

carina

195

sublateral
~~ process

194

Figs 191-195 Male genitalia of Acalyptris species, Belize. 191, laxibasis, holotype (29128 - BMNH), capsule; 192, same,
aedeagus; 193, species 29135 (29135 — BMNH), capsule; 194, same, valva; 195, same, aedeagus. Scale: 0.1 mm.

106 R. PUPLESIS AND G.S. ROBINSON

197

uncus
pseuduncus

198

pseuduncus

Figs 196-202 Male genitalia of Acalyptris species, Belize. 196, dividua, holotype (29125 - BMNH), capsule; 197, same,
paratype (AD0309 — VPU), cornutus; 198, same, holotype (29125 — BMNH), aedeagus; 199, platygnathos, holotype
(29132 — BMNH), uncus and pseuduncus; 200, same, aedeagus; 201, same, gnathos; 202, same, capsule. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 107

sublateral
process

cathrema

Figs 203-206 Male genitalia of Glaucolepis species. 203, aerifica, lectotype, Peru (28965 — BMNH), capsule; 204, same,
aedeagus; 205, argentosa, holotype, Belize (29106 — BMNH), aedeagus; 206, same, capsule. Scale: 0.1 mm.

108 R. PUPLESIS AND G.S. ROBINSON

+ vestibulum

[
Af
accessory sac

e
SS folded corpus
bursae

apophyses
anteriores

apophyses
posteriores

ductus
spermathecae or

ae ae corpus bursae
BOD eS” Bae NG reas
; ’ re
¥ = ce t
> Se ae ae 2 a
LS guaiatg ENE Gope we. oe
Se NT eo as ’
va r. , oF a
en , Tas ” rs eS annnn”

corpus bursae B ptadem BPE a o> fy
Ree ou? AP eine Cras

207

Figs 207,208 Female genitalia. 207, Manoneura basidactyla [non-type], Belize (29121 — BMNH); 208, Stigmella rudis,
paratype, Chile (DiSkus177 — ZMUC). Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 109

apophyses
posteriores

apophyses
anteriores

bursae

accessory sac

accessory sac

ductus ; /
spermathecae } j |

209 ioe

Figs 209, 210 Female genitalia of Stigmella species, Peru. 209, andina, paralectotype (28841 — BMNH); 210, marmorea,
paratype (Diskus181 — ZMUC). Scale: 0.1 mm.

R. PUPLESIS AND G.S. ROBINSON

110

212

214

Figs 211-214 Female genitalia of Stigmella species, Peru. 211, epicosma, paralectotype (28850 — BMNH); 212, epicosma,
non-type (Diskus 186 —ZMUC); 213, schoorli, paratype (Dikus206 — ZMUC); 214, paratype (Diskus 204 — ZMUC),

bursa. Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE 111

apophyses
posteriores

accessory sac

ust

le}

t

OUR er

Figs 215,216 Female genitalia of Stigmella species. 215, olyritis, paralectotype, Peru (28852 — BMNH); 216,
plumosetaeella, non-type, Mexico (Diskus 006 — USNM); 217, guittonae, paratype, Argentina (DiSkus2043 — MACN).

Scale: 0.1 mm.

, R. PUPLESIS AND G.S. ROBINSON

vestibulum

vestibulum

Om
eS eo
a

=

i en 3
five) Frese
& ry
af €
eG Wcccee td

Figs 218-220 Female genitalia, paratypes, Belize. 218, Stigmella pruinosa (29126 — BMNH); 219, Fomoria diskusi (28846
—BMNH); 220, Acalyptris bovicorneus (29116 — BMNH). Scale: 0.1 mm.

CENTRAL AND SOUTH AMERICAN NEPTICULIDAE

4
/ apophyses
iA anteriores
ae
?
X |
«|
y
5
is vaginal sclerite
<a .
Q)
2s
SN
aoa

s AO
[ é + ce
c

¢

AU ATTA

L
yi Z
~ ee OK v
ea

221

Figs 221,222 Female genitalia. 221, Acalyptris species 29140, Belize (29140 — BMNH); 22, Glaucolepis aerifica,
paralectotype, Peru (28966 — BMNH) [identity questionable — see text]. Scale: 0.1 mm.

113

114 R. PUPLESIS AND G.S. ROBINSON

Fig. 223 Distribution map of Nepticulidae species recorded from the Neotropical Region: 1, Enteucha cyanochlora; 2, E.
gilvafascia; 3, E. hilli; 4, E. contracolorea; 5, E. terricula; 6, E. snaddoni; 7, Manoneura basidactyla; 8, M. trinaria; 9,
Stigmella andina; 10, S. cuprata; 11, S. johannis; 12, S. rudis; 13, S. marmorea; 14, S. peruanica; 15, S. epicosma; 16, S.
schoorli; 17, S. hamata; 18, S. imperatoria; 19, S. olyritis; 20, S. eurydesma; 21, S. albilamina; 22, S. fuscilamina; 23, S.
gossypii; 24, S. kimae; 25, S. plumosetaeella; 26, S. barbata; 27, S. pruinosa; 28, S. ovata; 29, S. hylomaga; 30, S.
costalimai; 31, S. guittonae; 32, Ectoedemia reneella; 33, E. helenella; 34, E. mesoloba; 35, E. species 29105; 36, E.
fuscivittata; 37, Fomoria molybditis; 38, F. diskusi; 39, F. species 29122; 40, F- latipennata; 41, Acalyptris bovicorneus; 42,
A. martinheringi; 43, A. fortis; 44, A. hispidus; 45, A. novenarius; 46, A. lascuevella; 47, A. bifidus; 48, A. trifidus; 49, A.
tenuijuxtus; 50, A. unicornis; 51, A. laxibasis; 52, A. bicornutus; 53, A. species 29135; 54, A. dividua; 55, A. platygnathos;
56, A. species 29140; 57, Glaucolepis aerifica; 58, G. argentosa.

Bulletin of The Natural History Museum
Entomology Series

Earlier Entomology Bulletins are still in print. The following can be ordered from Intercept (address on inside front
cover). Where the complete backlist is not shown, this may also be obtained from the same address.

Volume 47

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A new genus of oriental lacewings (Neuroptera: Chrysopidae). S.J. Brooks. 1983. Pp. 1-26, 95 figs. £4.00

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Volume 52
No. 1 The sandflies of Egypt (Diptera: Phlebotominae). R.P. Lane. 1986. Pp. 1-35, 80 figs, 4 tables. £5.60
No. 2 Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae). G.S. Robinson. 1986. Pp. 37-181, 200
figs. £24.00
No. 3 A revision of the European Agathidinae (Hymenoptera: Braconidae). G.E.J. Nixon. 1986. Pp. 183-242, 68
figs. £11.00
No. 4 A key to the Afrotropical genera of Eucoilidae (Hymenoptera) with a revision of certain genera. J.

Quinlan. 1986. Pp. 243-366, 359 figs. £21.00

Volume 53

No. 1 A review of Miletini (Lepidoptera: Lycaenidae). J.N. Eliot. 1986. Pp. 1-105, 108 figs. £18.00
No. 2 Australian ichneumonids of the tribes Labenini and Poecilocryptini. ILD. Gauld & G.A. Holloway. 1986.
Pp. 107-149, 65 figs. £8.40
No. 3 The tribe Pseudophloeini (Hemiptera: Coreidae) in the Old World tropics with a discussion on the
distribution of the Pseudophloeinae. W.R. Dolling. 1986. Pp. 151-212, 121 figs. £11.50
No. 4 The songs of the western European grasshoppers of the genus Omocestus in relation to their taxonomy
(Orthoptera: Acrididae). D.R. Ragge. 1986. Pp. 213-249, 128 figs. £7.50
No. 5 The structure and affinities of the Hedyloidea: a new concept of the butterflies. M.J. Scoble. 1986. Pp.
251-286, 102 figs. £7.00
Volume 54
No. 1 Studies on the Old World species of Holothrips (Thysanoptera: Phlaeothripidae). S. Okajima. 1987. Pp. 1—
74, 207 figs. £14.00
No. 2 Spectacles and Silver Ys: a synthesis of the systematics, cladistics and biology of the Plusiinae (Lepidop-
tera: Noctuidae). I.J. Kitching. 1987. Pp. 75-261, 465 figs. £36.00
No. 3 A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr (Hymenoptera:
Formicidae). B. Bolton. 1987. Pp. 263-452, 100 figs. £36.50
Volume 55
No. 1 A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of certain
genera. M.W.R. de V. Graham. 1987. Pp. 1-392, 744 figs. £75.00
No. 2 The songs of the western European grasshoppers of the genus Stenobothrus in relation to their taxonomy
(Orthoptera: Acrididae). D.R. Ragge. 1987. Pp. 393-424, 116 figs. £6.00
Volume 56
No. | The legume-feeding psyllids (Homoptera) of the wet Palaearctic Region. I.D. Hodkinson & D. Hollis.
1987. Pp. 1-86, 294 figs. £16.00
No. 2 A review of the Malvales-feeding psyllid family Carsidaridae (Homoptera). D. Hollis. 1987. Pp. 87-127,
94 figs. £6.00
No. 3 A review of the Rhadalinae (=Aplocneminae) (Coleoptera: Melyridae). E.R. Peacock. 1987. Pp. 129-170,
59 figs. £8.00
No. 4 A revision of some Afrotropical genera of Eucoilidae (Hymenoptera). J. Quinlan. 1988. Pp. 171-229, 199
figs. £11.00
Volume 57
No. 1 A survey of the Ophioninae (Hymenoptera: Ichneumonidae) of tropical Mesoamerica with special
reference to the fauna of Costa Rica. I.D. Gauld. 1988. Pp. 1-309, 352 figs, 32 maps. £52.00
No. 2 A taxonomic revision of Alabagrus (Hymenoptera: Braconidae). M.J. Sharkey. 1988. Pp. 311-437, 28
figs, 22 maps. £24.50
No. 3 A taxonomic revision of Caryocolum (Lepidoptera: Gelechiidae). P. Huemer. 1988. Pp. 439-571, 221 figs.
£25.00
Volume 58

No. 1 The mealybug genus Planococcus (Homoptera: Pseudococcidae). J.M. Cox. 1989. Pp. 1-78, 40 figs.
No. 2 The Simuliidae (Diptera) of the Santiago onchocerciasis focus of Ecuador. A.J. Shelley, M. Arzube & C.A.
Couch. 1989. Pp. 79-130, 153 figs (including 2 plates in colour).

Volume 59

No. 1 The songs of the western European bush-crickets of the genus Platycleis in relation to their taxonomy
(Orthoptera: Tettigoniidae). D.R. Ragge. 1990. Pp. 1-35.
A reclassification of the Melanotus group of genera (Coleoptera: Elateridae). C.M.F. von Hayek. 1990.
Pp. 37-115.

No. 2 The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). S.J. Brooks & P.C.
Barnard. 1990. Pp. 117-286.

Volume 60
No. 1 The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). P.H. Williams. 1991.
Pp. 1-204.

No. 2 Sattleria: a European genus of brachypterous alpine moths (Lepidoptera: Gelechiidae). L.M. Pitkin & K.
Sattler. 1991. Pp. 205-241.
A review of wing reduction in Lepidoptera. K. Sattler. 1991. Pp. 243-288.

Volume 61

No. | Thrips (Thysanoptera) from Pakistan to the Pacific: a review. J.M. Palmer. 1992. Pp. 1-76.

No. 2 Neotropical red-brown Ennominae in the genera Thysanopyga Herrich-Scaffer and Perissopteryx Warren
(Lepidoptera: Geometridae). M. Kruger & M.J. Scoble. 1992. Pp. 77-148.

Volume 62

No. | Caloptilia leaf-miner moths (Gracillariidae) of South-East Asia. Decheng Yuan and Gaden S. Robinson.
1993. Pp. 1-37.

No. 2 Neotropical Emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular
reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Linda M. Pitkin. 1993.
Pp. 39-159.

Volume 63

No. 1 A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae), parasitoids of the

immature stages of economically important insect species (mainly Hemiptera and Lepidoptera). D.W.
Huang and J.S. Noyes. Pp. 1-135.

No. 2 A taxonomic review of the common green lacewing genus Chrysoperla (Neuroptera: Chrysopidae). S.J.
Brooks. Pp. 137-210.

Volume 64

No. 1 Revision of the neotropical genus Oospila Warren (Lepidoptera: Geometridae) M.A. Cook and M.J.
Scoble. Pp. 1-115.

No. 2 Encyrtidae of Costa Rica (Hymenhoptera: Chalcidoidea): the genus Aenasius Walker, parasitoids of
mealybugs (Homoptera: Pseudococcidae). J.S. Noyes and H. Ren. Pp. 117-164.

Volume 65

No. 1 A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae).
Y. Zhang and M.D. Webb. Pp 1-103.

No. 2 Encyrtidae (Hymenoptera: Chalcidoidea) of Costa Rica: the genera and species associated with jumping

plant-lice (Homoptera: Psylloidea). J.S. Noyes and P. Hanson. Pp. 105-164.

Volume 66

No. 1 Biosystematic studies on the Simuliidae (Diptera) of the Amazonia onchocerciasis focus. A.J. Shelley,
C.A. Lowry, M. Maia-Herzog, A.P.A. Luna Dias and M.A.P. Moraes. Pp. 121.

No. 2 Microtermes in East Africa (Isoptera: Termitidae: Macrotermitinae) S. Bacchus. 1997. Pp. 123-171.

Volume 67

No. | Mealybugs of the genera Eumyrmococcus Silvestri and Xenococcus Silvestri associated with the ant genus
Acropyga Roger and a review of the subfamily Rhizoecinae (Hemiptera, Coccoidea, Pseudococcidae). D.J.
Williams. 1998. Pp. 1-64.
Monophyly of the dacetonine tribe-group and its component tribes (Hymenoptera: Formicidae). B. Bolton.
1998. Pp. 65-78.
An annotated checklist of bumble bees with an analysis of patterns of description (Hymenoptera: Apidae,
Bombini). P.H. Williams. 1998. Pp. 79-152.

No. 2 A revision of the leafhopper tribe Paraboloponini (Hemiptera: Cicadellidae: Selenocephalinae) in the
Indian subcontinent. C.A. Viraktamath. 1998. Pp. 153-208.
A review of the Palaearctic Neorhacodinae (Hymenoptera, Ichneumonidae) with Eremura Kasparyan,
1995 new to the west Palaearctic. D.G. Notton and M.R. Shaw. 1998. Pp. 209-218.

Volume 68

No. 1 Revision of the Oriental Opostegidae (Lepidoptera) with general comments on phylogeny within the
family. Rimantas Puplesis and Gaden S. Robinson. 1999. Pp. 1-92.

No. 2 A revision of the African and Malagasy species of the genus Leptomastix (Hymenoptera, Encyrtidae),
parasitoids of mealybugs (Homoptera: Pseudococcidae). Jean-Marc Anga and John S. Noyes. 1999. Pp.
93-128.
A revision of the north-west European species of the formosus species group of Spilomicrus (Hymenop-
tera: Diapriidae). David G. Notton. 1999. Pp. 129-144.
A review of the soldierless African termite genus Amicotermes Sands 1972 (Isoptera, Termitidae,
Apicotermitinae). W.A. Sands. 1999. Pp. 145-193.

CONTENTS

Editorial
Ri Vane-Wright
_Areview of the Central and South American Nepticulidae
with special reference to Belize
Rimantas Puplesis and Gaden S. Robinson

ENTOMOLOGY SERIES

Vol. 69, No. 1 , June 2000