Revista Brasileira de Entomologia

http://dx.doi.org/10.1590/S0085-56262013005000012

A new genus and species of leaf miner (Lepidoptera, Gracillariidae) for

Chile associated to the native tree Lithraea caustica
Enrique A. Mundaca1, Luis E. Parra2 & Hctor A. Vargas3
1

Universidad Catlica del Maule, Facultad de Ciencias Agrarias y Forestales, Escuela de Agronoma, Casilla 7-D, Curic, Chile. [email protected]
Departamento de Zoologa, Facultad de Ciencias Naturales y Oceanogrficas, Universidad de Concepcin. Casilla 160-C, Concepcin, Chile.
[email protected]
3
Departamento de Recursos Ambientales, Facultad de Ciencias Agronmicas, Universidad de Tarapac, Casilla 6-D. Arica, Chile. [email protected]
2

ABSTRACT. A new genus and species of leaf miner (Lepidoptera, Gracillariidae) for Chile associated to the native tree Lithraea
caustica. We propose the new genus and species of Gracillariidae (Lepidoptera) Hualpenia lithraeophaga Mundaca, Parra &Vargas
gen. nov., sp. nov., leaf miner of Lithraea caustica (Mol.) H. et Arn (Anacardiaceae) occurring in southern central Chile. Aspects of the
life cycle, adult and larval morphology, development and feeding habits of the new genus and species are also presented. We emphasise
the uniqueness and importance of this new species for broadening the current knowledge on the Chilean fauna of Gracillariidae.
KEYWORDS. Hualpn, hypermetamorphic, Insecta.

The family Gracillariidae includes small-sized moths with

white marks on the wings and is considered the main group
of lepidopteran leaf-miners in the world (Davis 1987). Most
species of this family have a remarkable larval development
characterised by having at least two well-differentiated morphological stages, although authors such as Trgardh (1913)
recognise up to four different stages. This type of development, with differentiated larval stages, has been called hypermetamorphosis or heteromorphic development (Hinton
& Mackerras 1970; Kumata 1978; Wagner et al. 2000). The
first stages are normally a flat-shaped larva with ambulatory
callosities instead of properly developed legs, simplified
chaetotaxy and highly modified mandibles. These specialised
mandibles are generally scissor-like shaped and serve to
saw the plant tissue, separating the cuticle from the epidermis. These early stages are characteristically sap feeding. The
second developmental stages have the typical eruciformshaped caterpillar, typical of the order Lepidoptera, with
conspicuous legs and prolegs, a spinneret and well-developed chewing mandibles to feed on tissues. This larval stage
has been called a tissue feeding form (Trgardh 1913).
The latest checklist of Neotropical Gracillariidae indicated a total of 179 species distributed in this region (De
Prins & De Prins 2012). Some of these species have been
described in recent taxonomic contributions (e.g. Davis &
Wagner 2005, 2011; Vargas & Landry 2005; Vargas & Parra
2005; Landry 2006; Kawahara et al. 2009). However, further
additional fieldwork is required to obtain better records and
to increase the understanding of the evolutionary history of
this interesting fauna.
The Chilean native fauna of Gracillariidae is relatively
unknown, with only five native species described so far (De
Prins & De Prins 2012): Parectopa rotigera Meyrick, 1931,

Prophyllocnistis epidrimys Davis, 1994, whose larvae are leaf

miners of the native plant Drimys winteri J.R. et G. Forst,
1776 (canelo); Phyllocnistis puyehuensis Davis, 1994;
Chileoptilia yaroella Vargas & Landry, 2005; and Angelabella
tecomae Vargas & Parra, 2005. The introduced pest species
Phyllocnistis citrella Stainton, 1856 is restricted to the northern coastal valleys associated with citrus plantations.
The VIII Region of Chile, which includes the Hualpn
Peninsula (Fig. 1), is a transitional area where two different
forest communities overlap and become in contact (Polymeris
1995). These two types of communities are the sclerophyllous
forest, part of the Mediterranean zone (which comes from
the north), and the hygrophyllous forest, a vegetation type
that characterises the sub-Antarctic area. In this transitional
area, the vegetation acquires a particular pattern, which is
different from the two main zones. The dominant tree species are: Cryptocarya alba (Mol.) Looser, 1782 (peumo),
Aextoxicon punctatum Ruiz & Pav (olivillo), L. caustica (litre) and Peumus boldus Molina, 1782 (boldo). The Hualpn
forest should be considered the northern limit of the Valdivian
forest, although the main characteristics of the forest remain
mainly sclerophyllous. There are 19 different vegetation types
described for Hualpn Peninsula by Polymeris (1995) that
include a wide variety of different habitats, ranging from
grasslands and artificial plantations to secondary growth
patches of fragmented native forest and shrubs. Lithraea
caustica (Mol.) H. et Arn has been described as the dominant species of at least 5 vegetation types described for the
peninsula (Polymeris 1995). In addition, L. caustica is a shrub
that belongs to the Anacardiaceae family, whose species are
typical of the Matorral Maulino (Maulean thicket), which is
dominated in its higher strata by sclerophyllous species
(Hoffman 1982). The species of this family are characterised
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Mundaca et al.

by having thick, resin-like sap juices which are frequently

caustic or poisonous (Cabrera 1938) and are known for provoking allergic reactions on humans.
The aim of this study is to describe and provide biological background information of a new genus and species of
Gracillariidae for the Chilean fauna of leaf miners associated with the native tree Lithraea caustica.
METHODS
Sampling was carried out in the Hualpn Peninsula,
3645S7313O (Fig. 1) between March and November,
1997. We collected infected leaves by inspecting L. caustica
trees in the field. Leaves containing larvae were collected
from the plant and kept in plastic bags to be transported to
the laboratory. Larvae utilised for character description were
fixed in alcohol 70%. A number of larvae were kept alive
inside the leaves in order to rear them into adults. Damages
on the leaves were photographed to show the type of damage
provoked by the different larval stages. Electronic microscope
pictures were taken in the Scanning Electron Microscope
(SEM) laboratory of the Universidad de Concepcin, Chile.
Examined material is deposited in the Museo de Zoologa
Universidad de Concepcin (MZUC), Concepcin, Chile. The
female genitalia is not illustrated since the slide containing
its preparation was lost during the 2010 earthquake in Chile.
TAXONOMY
Hualpenia Mundaca, Parra & Vargas gen. nov.
Type species: Hualpenia lithraeophaga Mundaca, Parra & Vargas sp. nov.

Diagnosis. Among the Neotropical genera, Hualpenia is

similar to Eucosmophora Walsingham, 1897, by the presence
of a digitate lobe on the costal margin of the valva in the male
genitalia. However, the veins R5 and M1 are separate on the
forewings of Eucosmophora, meanwhile these veins are stalked
in Hualpenia. Veins R5 and M1 are also separate in the forewings of other non-Neotropical genera with similar digitate lobe
on the costal margin of the valve: Amblyptila Vri, 1961,
Borboryctis Kumata & Kuroko, 1988, Gibbovalva Kumata &
Kuroko, 1988, Melanocercops Kumata & Kuroko, 1988,
Phodoryctis Kumata & Kuroko, 1988, and Sauterina
Kuznetsov, 1979. Among the Neotropical genera, veins R5 and
M1 are stalked in Chileoptilia Vargas & Landry, 2005; however, the morphology of the male genitalia is remarkably different, lacking a digitate lobe on the costal margin of the valve.
The same is true when comparing Hualpenia with other nonNeotropical genera with stalked R5 and M1.
Description. Small moths with 1011 mm forewing length
(Fig. 2). Forewings colour brown cooper, lanceolated, with
white marks on the distal three-quarters. Hindwings fringed,
grey coloured, shorter than the forewings. Forewing venation
with all veins present (Fig. 5A). Base of R3, R4, R5+M1 and
M2 not differentiated; distal margin of discal cell slightly differentiated. R5 extended towards apex; A1+2 and CuP weak,
Revista Brasileira de Entomologia 57(2): 157164, June 2013

Fig. 1. Hualpn Peninsula map.

poorly marked; M1 fused with R5. Hindwing venation with

Sc+R1, R5 and Cu2 present. Male genitalia with a digitate lobe
in the costal margin of the valve. Hypermetamorphic larva
with leaf miner habits. First larval instars flat shaped, sap feeders, forming serpentine mines (Fig. 3A), with ambulatory callosities instead of legs and prolegs. Later instars, larva
eruciform tissue-feeding forming patchy mines (Fig. 3B). Last
instar larva changes its colour from pale white-grey to dark
pink prior to pupation. The cocoon remains inside the mine,
surrounded by a silk chamber until the adult emerges.
Hualpenia lithraeophaga Mundaca, Parra & Vargas sp.nov.
Description. Male. Head. Head roughly short haired (Fig.
4A).With a tuft of hairs, directed forward and coming out from
the lateral margins of the forehead, under the antennae. Face
smooth, covered by white scales. Labial palpus covered by
white scales which become dark brown towards the last two
segments. Ocelli absent. Haustellum covered with setae (Fig.
4B). Maxillary palpus covered with scales, shorter than distal
segment of labial palpus. Distal segment of the labial palpus
same length of the compound eyes diameter. Filiform antennae. Pillifers present. Thorax. Light brown coloured. Ventral
area dark brown with two bands of white scales covering pro
and mesothorax long each side. Legs long and slender. Forewing brown-silver lanceolated, apical region dark cooper-ferruginous with a white semicircular band on the distal threequarters; a smaller white mark also present, closer to the apical region of the wing. Distal border of the wing black coloured.
Hindwings grey-silver, lighter, shorter than the forewings with
a uniform colour pattern (Fig. 2). Female with a frenulum con-

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159

1 mm

3A

3B

Figs. 23. 2. Hualpenia lithraeophaga sp. nov., holotype male adult, 11 mm wingspan. 3. Leaf damage provoked by sap-feeding larva (A) and by tissue
feeding larva (B) of Hualpenia lithraeophaga sp. nov.

sisting of two setae. Male with frenulum consisting of only

one setae. Distal portion of the hindwings fringed. Abdomen.
Dorsal area dark brown coloured; ventral area light brown to
yellowish; A8 membranous, covered with a ventral band of
white scales. Genitalia. Dorsal arms of the saccus V shaped.
Valve four times longer than wide, hairy towards apex. Medial process slightly bent. Short, digitate lobe arising near 1/4
costa. Saccus subtriangular, rounded towards apex. Uncus and
transtilla absent. Tegumen subtriangular enlarged, weakly sclerosed. Coremmata absent. Aedeagus long, same length than
valve. No cornuti (Fig. 5B).
Female. Similar to male, but abdomen with ventral area
grey silver, without the band of white scales on the eight
segment. Papillae analis covered by long setae (Fig. 4C).
Type Material. Holotype male (Fig. 2), December 7, 1999 (MZUC);
Allotype female, November 25, 1999 (MZUC). Paratype: 1 female, November 25, 1999 (used for the electron microscope preparation). Holotype,
allotype and paratype were reared in the laboratory from larvae collected at
Parque Botnico (Universidad de Concepcin), in Hualpn Peninsula, VIII
Regin, Chile. Enrique Mundaca leg. Material deposited at MZUC.

Additional Examined material. One male, December 7, 1998; two females reared on January 14, 1999. One of the females was utilised to describe wing venation and genitalia preparation. A second female specimen
was used for the Scanning Electron Microscope (SEM) analysis. All specimens obtained from larvae collected by Enrique Mundaca at the type locality. Material deposited at MZUC.

Sap-feeding instar larva. Leaf miner of Lithraea caustica.

Head light brown coloured. Body white to light brown, with
plates of micro spines on the dorsal and ventral region of
segments T1T3 and A1A9 (Fig. 6A, 6C and 6D). Body
and cephalic capsule depressed (Fig. 6A, 7A and 7B). Ambulatory callosities present in thorax and abdomen (Fig. 6E
and 7B). Scissor-like mandibles, horizontally arranged (Fig.
6F) with irregular saw-like borders (Fig. 6B). Spinneret not
observed. Reduced chaetotaxy.
Tissue feeding larva. Head. Hypognatus head (Fig. 8A).
Spinneret present (Fig. 8B). Ocular area with 4 distinct stemmata (Fig. 8C). Setae A1, A2 and A3 present. A1 to the same
distance from stemma4 than A2 from stemmata 2 and 3. A3
located half a distance from stemma1. SS2 laterally located
behind stemma1 in front of stemma2. SS3 ventrally located,
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Fig. 4. Hualpenia lithraeophaga sp. nov. (A) Adult head, ventral-front view. (B) Proboscis. (C) Papillae analis covered by setae.

100x

Fig. 5. Hualpenia lithraeophaga sp. nov. (A) Wing venation. (B) Male genitalia ventral view, and (C) aedeagus left view.

Fig. 6. Hualpenia lithraeophaga sp. nov. (A) Dorsal view of sap-feeding larva. (B) Details of the scissor-like mandible in the sap-feeding larva. (C) Sapfeeding larva, dorsal view of the head capsule. (D) Dorsal view of plates covered with micro spines present on thoracic segments of the sap-feeding larva.
(E) Lateral view of partially developed legs. (F) Lateral view of the scissor-like mandibles present on the sap-feeding larva.
Revista Brasileira de Entomologia 57(2): 157164, June 2013

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161

1 mm

1 mm

Fig. 7. Hualpenia lithraeophaga sp. nov. (A) Dorsal view of sap-feeding larva. (B) Lateral view of sap-feeding larva. (C) Tissue feeding larva. (D) Lateral
view of tissue feeding larva.

Fig. 8. Hualpenia lithraeophaga sp. nov. (A) Dorsal view of the head of the tissue-feeding larva. (B) Tissue-feeding larva, maxillae and labium. (C) Detail
of the ocular area of the tissue-feeding larva. (D) Frontal detail of the head of the tissue-feeding larva. (E) Lateral view of tissue-feeding larva legs. (F)
Ventral view of prolegs. (G) Details of the crochets.

surrounded bystemma4. L1 present, shorter than A3. C1 and

C2 present on the Clypeus. C1 one third the length of C2 (Fig.
8D). Thorax. Prothorax. D1 present on the cervical shield.
SD1, SD2 and D2 absent. Complex L1 and L2 present. L1
located above the spiracle and closer to L2 than D1. L2 located under the spiracle. SV1 present, twice longer than L1
and L2 (Fig. 9A). Mesothorax and metathorax. D1, D2, SD1
and SD2 on the mesothoracic shield. L1 same length than L2.
SV1 present, 1.5 times longer than L1 and L2. Spiracle absent. Legs well developed (Fig. 10B, 11B, 21 and 22). Abdomen. Spiracles absent in A9 and A10. D1, D2 and SD1 present
in all other abdominal segments (A18). SD2 present only in
A3. SD3 present only in A10. L1 occurring in all segments,
always ventrally oriented to the spiracle.L2 three quarters the
length of L1, present in A1 and A2.SV1and SV2 absent from
A10. SV3 absents from A1, A9 and A10. V1 present from A3

to A6 (Fig. 9A). Prolegs with only one row of uniordinal crochets arranged in an uniordinal manner following criteria established by Stehr (1987) (Fig. 8F and 8G).
Examined material. One larva, collected on April 15, 1997, 5 larvae
collected on May 28, 1997, 1 larva collected on June 27 1997, 2 larvae
collected on September 26, 1997. All specimens collected at Parque
Botnico (Universidad de Concepcin) Hualpn Peninsula, VIII Regin,
Chile, Enrique Mundaca leg. Material deposited at MZUC.

Pupa. Obtecta type. Maximum length 5 mm. Maximum

diameter 1mm. Integument dark brown. Antennae about 1/5
longer than the body. Frontal process (cocoon cutter) short,
slightly pointed towards apex. Labial palpi visible. Galea
shorter than forewings. Forewings extending almost to A7,
parallel, not sealed in front of the body. Middle and hind legs
visible, partially covered by the forewings (Fig. 9B).
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Mundaca et al.

T1

T2

A1

A2

A3

1 mm

A4

A6

A9

A10

Fig. 9. Hualpenia lithraeophaga sp. nov. (A) Chaetotaxy of thoracic segments: 12; and abdominal segments: 1, 2, 3, 4, 6, 9 and 10. (B) Pupa, lateral,
ventral and dorsal view.

Examined material. One pupa collected on August 1989, at Parque

Botnico (Universidad de Concepcin) Hualpn Peninsula, VIII Region
Chile, Hector Ibarra leg. Material deposited at MZUC.

Egg. Unknown.
BIOLOGY
So far, it has not been possible to collect eggs in the field.
Marks left on the leaves and the number of larvae found in
the leaves show that there is just one egg laid on the adaxial
surface of the leaf. Only in one occasion we have observed a
mine in the abaxial surface. The larva inside was dead and at
the sap-feeding instar.
Body structure of the last larval instar differs from other
species of the family, such as those of the genera Marmara
Clemens, 1863, (Fitzgerald & Simeone 1971) and Prophyllocnictis Davis, 1994, where the second developmental stage
does not feed and belongs to the spinning type (Kumata 1978)
and resembles a typical eruciform lepidopteran larva. Prior
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to the pupation, the larva suffers a remarkable change of its

colour, from pale grey to dark pink. This phenomenon has
been observed during late winter-early spring season, at the
beginning of September. Pupation occurs inside the mine,
although this observation was carried out in controlled laboratory conditions. No cocoons or pupae have been observed
inside the leaf mine in the field.
Phenology. Our observations in the field indicate that
the life cycle of this species is univoltine, with adults flying
during late spring and early summer (Table I). Although it is
possible to observe mines throughout the year, many of them
were observed with dead larvae inside the galleries. Dead
larvae are easy to identify because of the colour of the mines
as they become pale grey. Table I shows the records of different stages of the species and the month that have been
recorded to be occurring in the field. These results are based
on the observation of living larvae inside the mines.
Leaf damage pattern. There are basically two types of
mines produced by the larvae. Serpentine mines (Fig. 3A) are

A new genus and species of leaf miner for Chile associated to the native tree Lithraea caustica

Table I. Phenology of Hualpenia lithraeophaga Mundaca, Parra & Vargas.

Stages
Egg
Larva
Pupa
Adult

Months
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
?
?
?
X
X
X X X
X
X
X X
?
?
X X

produced by the sap-feeding instars. This type of leaf mine is

pale coloured and characterised for not having the cuticle too
separated from the parenchyma. At this stage the mine looks
narrow, whitish in colour, serpentine-shaped and flat. The second type of mine has a patch shape (Fig. 3B). It occurs after
the hypermetamorphosis of the larva, when the larva changes
its feeding habits and morphs to a tissue-feeding larva type.
At this stage the mine looks externally brownish, with the leaf
cuticle clearly separated from the parenchyma giving the leaf
a swollen appearance. It is common to observe in the field
leaves with the cuticle broken without the larva inside. When
examining the interior of the mine, it is possible to find rests
of head capsules and droppings of the larvae.
DISCUSSION
The characteristics of this new genus allow locating it in
the subfamily Gracillariinae. The most important characters
described by Davis & Robinson (1999), as representative of
Gracillariinae, are present in this species: adults raised at
steep angle while perching, Rs archers towards Sc, eighth
abdominal segment membranous in the male, and A9 with
bisetose (D1D2) complex in the tissue-feeding larva. We
compared head, thorax and male genitalia characters of this
genus with those described for the type species of genera of
Gracillariinae proposed by Davis & Robinson (1999) and
described by Vari (1961). The head-scales arrangement in
Hualpenia is more similar to Epicephala Meyrick, 1880, and
Aristaea Meyrick, 1907, but it differs from Aristaea in that it
does not have ocelli. Maxillar and labial palpi seem to follow the pattern proposed for most of the representative genera of Gracillariinae, where labial palpi are longer and more
conspicuous than the maxillary palpi. Male genitalia of
Hualpenia, compared to that of other genera of Gracillariinae,
does not show the presence of coremmata. However, this
character does not exclusively belong to Hualpenia, as for at
least in three other genera of Gracillariinae, namely,
Eucosmophora Walsingham, 1897, and Schedocercops Vri,
1961, the presence of coremmata was not reported.
This genus differs from most Gracillariinae in that the
last instar changes its colour, from pale white-grey to dark
pink prior to pupation. Another remarkable character of
Hualpenia is the presence of a ventral band of white scales
covering the males 8th abdominal segment which is not
present in the females abdomen. Thus, Hualpenia can be
defined mainly by a combination of characters, which cannot be found in other genera of Gracillariinae.

163

Hualpenia lithraeophaga is the first Chilean Gracillariidae

with larvae associated with Anacardiaceae. Previous host plant
record for the Chilean fauna were restricted to the plant families Bignoniaceae (Vargas & Parra 2005), Fabaceae (Vargas &
Landry 2005), Malvaceae (Vargas et al. 2013) and Winteraceae
(Davis 1994). Host plant association with Anacardiaceae has
been previously mentioned for other three species of South
American Gracillariidae in the genera Caloptilia Hbner, 1825,
Eucosmophora Walsingham, 1897, and Leurocephala Davis
& McKay, 2011 (Davis et al. 2011), with most surveys for
Gracillariidae on South American Anacardiacaeae directed to
the Brazilian Pepper Tree, Schinus terebinthifolius Radii, 1820.
The genus Lithraea has been also reported as host plant for
two of these species: Caloptilia schinusifolia Davis & Wheeler,
2011, and Eucosmophora schinusivora Davis & Wheeler, 2011
(Davis et al. 2011).
The finding of this remarkable species enhances the necessity for conserving the last remnants of native vegetation
on the Hualpn Peninsula. As mentioned before, this is the
last relict of transitional schlerophyllous-latifoliated evergreen forest of Chile (Gajardo 1994; Polymeris 1995) and
is considered also a unique ecosystem in the world. The
Hualpn Peninsula is currently under an accelerated process
of change, mainly due to economic activities (Martinez &
Donoso 1997) such as urbanisation, construction of recreational spaces and forestry. Such changes do not consider
important the protection of the natural uniqueness of an area
that could be the potential refuge of a number of unknown
species.
ACKNOWLEDGEMENTS
We would like to thank all those people that have helped
us to carry out this research: to Dr. Viviane Jerez, Dr. Thomas Ogden, Minerva Contreras, Patricia Bocaz, Romina
Villagrn and Mara de Los ngeles, many thanks for your
help in our field trips and laboratory work. To Hiplito Rifo
and Gabriel Mendoza for preparing the adult and larvae figures. To Mario Moya for edition of the electronic figures. We
would also like to thank the personnel in charge of the Scanning Electron Microscope for their help during the preparation of the samples and Mariana Lazzaro-Salazar for her help
with the figures edition.
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Received 8 November 2012; accepted 4 January 2013

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