Systematic Entomology (2013), 38, 334–348

A molecular analysis of the Gelechiidae (Lepidoptera,

Gelechioidea) with an interpretative grouping of its taxa
1 2 3
O L E K A R S H O L T , M A R K O M U T A N E N , S A N G M I L E E and L A U R I
4
KAILA
1
Zoological Museum, The Natural History Museum of Denmark, Copenhagen, Denmark, 2 Department of Biology, University of
Oulu, Oulu, Finland, 3 School of Life Sciences, Arizona State University, Tempe, AZ, U.S.A., and 4 Finnish Museum of Natural
History, Zoology Unit, University of Helsinki, Helsinki, Finland

Abstract. We re-examine the higher level phylogeny and evolutionary affinities of

the family Gelechiidae (Lepidoptera: Gelechioidea) based on DNA sequence data for
one mitochondrial gene (cytochrome c oxidase subunit I ) and seven nuclear genes
(Elongation Factor-1α, wingless, Ribosomal protein S5, Isocitrate dehydrogenase,
Cytosolic malate dehydrogenase, Glyceraldehyde-3-phosphate dehydrogenase and
Carbamoylphosphate synthase domain protein). Fifty-two taxa representing nearly
all established subfamilies and tribes of Gelechiidae, and about 10% of described
gelechiid genera, in addition to five outgroup taxa were sequenced. Data matrices
(6157 bp total) were analysed under model-based evolutionary methods (Maximum
Likelihood and Bayesian Inference), resulting in novel high-level phylogenetic
interrelationships. The best supported cladogram divided the Gelechiidae into six
distinct clades corresponding to the subfamilies Anacampsinae, Dichomeridinae,
Apatetrinae, Thiotrichinae, Anomologinae and Gelechiinae (+ Physoptilinae, which
were not available for study). The results suggest the following adjustments in
gelechiid interrelationships: Brachmini is nested within Dichomeridinae; Anarsiini
is the sister group of Chelariini; Pexicopiinae is the sister group of Apatetrinae,
here suggested to be treated as a tribe Pexicopiini of Apatetrinae. A new subfamily
Thiotrichinae (subfam.n.) is proposed on the basis of the resurrected genus Thiotricha
Meyrick (gen.rev.), which includes Macrenches Meyrick, Palumbina Rondani and
Polyhymno Chambers. Gelechiidae display a wide array of life-history strategies, but
the diversity in patterns of larval mode of life has direct phylogenetic correlation only
below subfamily level, suggesting multiple origins and/or frequent reversals for traits
such as external or internal feeding and leaf mining within the family.

Introduction and is among the most diverse Lepidoptera fauna in many

The micro-moth family Gelechiidae comprises c. 4700
described species in about 500 genera (Nieukerken et al.,
2011), with many species – probably at least equally many –
still to be described. The family has a worldwide distribution
Correspondence: Ole Karsholt, Zoological Museum, The Natural
History of Denmark, Copenhagen, Denmark.
E-mail: okarsholt@snm.ku.dk
regions and habitats. The expected increase in the number
of actual extant species is based not only on estimations
of already known but not formally described species (e.g.
Edwards, 1996), but also based on general difficulties in dis-
covery and recognition of species in insufficiently explored
tropical areas (Novotný & Basset, 2000; Janzen et al., 2005).
Moreover, cryptic diversity in apparently better known faunas
remains to be discovered (e.g. Nason et al., 2002; Huemer &
Karsholt, 2011).
Gelechiids are small- to medium-sized, often grey or brown
moths, whose larvae exhibit a wide range of feeding strategies;
feeding as leaf-miners, gall-inducers or stem-borers, or feeding

334 © 2013 The Royal Entomological Society


within flowers or fruits, between tied or rolled leaves, or
concealed within silken tunnels in the earth or within a larval
case. Their food includes almost all kinds of plants: herbs,
grasses, deciduous and coniferous trees, and also, mosses, ferns
and even lichens (Powell, 1980).
Larvae of Gelechiidae include a number of more or less
serious pests damaging crops or stored food products, causing
high economic losses. Zhang (1994) lists 258 species in 85
genera as being economically important. These include well-
known species such as Pectinophora gossypiella (Saunders)
(pink bollworm), Sitotroga cerealella (Olivier) (angoumois
grain moth), Phthorimaea operculella (Zeller) (potato tuber
moth) and Tuta absoluta (Meyrick) (tomato leaf miner), all
with an enormous printed and internet literature. For example,
a search on Google for T. absoluta on 3 March 2012 gave
445,000 links.
Hovever, gelechiids include potentially beneficial insects as
well as harmful ones. Larvae of Gelechiidae are generally
monophagous on their host plant species or genus, and several
species have been used as biological control agents against
invasive weeds around the world (e.g. Diatloff & Palmer, 1988;
Boggs et al., 1991; Klinken et al., 2003). The close connection
to a single or a few host plants, often correlated to strict
ecological demands, make gelechiids useful in environmental
studies and as indicators for biological conservation (e.g.
Björklund, 2005). Moreover, the entire fauna of Gelechiidae
has been evaluated for the purpose of red-listing in some
countries (e.g. Karsholt, 2006).
A closer scrutiny of Gelechiidae would undoubtedly reveal
an array of intriguing features; for example, species with
twirling (Common, 1990) or even dancing adults have been
described (Kawahara & Adamski, 2006). A few species
in Dichomeris, e.g. D. nonstrigella (Chambers), have adults
with very similar behaviour and posture as the cantharid
beetle Lucidota atra (Olivier); the latter being unpalatable
to visual predators including birds (J.-F. Landry, unpublished
data). The widespread Chrysoesthia drurella (Fabricius) differs
from its congeners by its red-orange forewings with silvery
and black markings, resembling species of other families
(e.g. Momphidae, Oecophoridae, Heliodinidae), and it is
likely that its striking appearance is part of a defensive
strategy, as these moths resemble red and black Heteroptera
(O. Karsholt, unpublished data). An intriguing phenomenon
recently discovered in the gelechiid genus Deltophora is the
ability of the adults of certain species to chemically break
up plant pollen, which is otherwise unknown in animals (Luo
et al., 2011).
A well-supported, phylogeny-based higher-level classifica-
tion for the major lineages of gelechiid moths has not been
published so far. Instead, there have been several contradic-
tory classifications suggested, none of which is based on a
really comprehensive phylogenetic evidence. Such a classifi-
cation would thus be of significance for the systematics of
this taxon, as well as for understanding the bionomics of pest
species and other intriguing traits.
The world fauna of Gelechiidae was last reviewed by
Meyrick (1925) and, based on that work, subsequently
© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348
Molecular analysis of Gelechiidae 335
catalogued by Gaede (1937). The study of this diverse family
is greatly hampered by the lack of a modern catalogue or
checklist. Meyrick (1925) proposed a classification of the
Gelechiidae – given as ‘Gelechiadae’ – based primarily on
the wing venation and to a lesser extent on features of
the labial palpi. He divided the family into nine genus-
groups (each named ‘type’): Apatetris, Aristotelia, Gelechia,
Anacampsis, Protolechia, Chelaria, Dichomeris, Lecithocera
and Autosticha, of which the two last mentioned mostly include
species now placed in the Autostichidae and Lecithoceridae.
Meyrick’s genus groups were later elevated to subfamilies by
Le Marchand (1947). They include: Apatetrinae, Aristoteli-
inae, Gelechiinae, Anacampsinae, Chelariinae, Dichomeridi-
nae, Lecithocerinae and Autostichinae. Le Marchand (1947)
did not erect a subfamily for the Australian Protolechia group,
and the systematic position of this group has remained dubious.
Meyrick did not consider characters of the genitalia in his
classification of the Gelechiidae. During the twentieth century
the genitalia became increasingly important in taxonomic
studies of Gelechiidae, both at the species and genus level,
especially initiated by Pierce & Metcalfe (1935) and Busck
(1939). Among more recent publications that emphasize the
value of genitalia are those of Sattler (1960, 1976), Povolný
(1964), Wolff (1958) and Huemer (1988). These and other
similar studies have focused at the generic, tribal or subfamilial
level of inquiry in specific geographical areas. Outstanding in
this respect is the work by Janse (1949–1963), who revised
the rich gelechiid fauna of southern Africa using both genitalia
and wing venation. However, Janse did not propose a higher
classification of the Gelechiidae to synthesize his results.
Meyrick’s (1925) old system has been modified by many
smaller and larger taxonomic works on Gelechiidae, e.g.
Hodges (1978) Becker (1984), Karsholt & Riedl (1996)
and Edwards et al. (1996). However, none of these revised
classifications included modern phylogenetic methods for
resolving the gelechiid subfamilies and tribes.
Hodges (1978, 1986, 1998) proposed different classifications
of the Gelechioidea, within which he elaborated on the
higher classification of the Gelechiidae. In the first of
these he proposed seven subfamilies of the Gelechiidae:
Anomologinae, Gelechiinae, Anacampsinae, Dichomeridinae,
Chelariinae, Lecithocerinae and Physoptilinae (Hodges, 1978).
Later (Hodges, 1986) he restricted the number of subfamilies
to three: Gelechiinae, Dichomeridinae and Pexiocopiinae,
mainly based on the structures of the abdominal support
system of sternum II, and transferred the Lecithocerinae and
Physoptilinae from the Gelechiidae. Finally, Hodges (1998)
used cladistic methods for the first time to construct a
phylogeny for the Gelechioidea, resulting in the inclusion
of the Physoptilinae in the Gelechiidae along with the three
subfamilies (Gelechiinae, Dichomeridinae and Pexiocopiinae)
which he recognized previously (Hodges, 1986).
Kuznetzov & Stelkov’nikov (1984, 2001) proposed an alter-
native classification based mainly on musculature morphol-
ogy of the male genitalia, and divided the Gelechiidae into
six subfamilies: Metzneriinae, Gelechiinae, Anacampsinae,
Teleiodinae, Chelariinae and Dichomeridinae. Ponomarenko
336 O. Karsholt et al.
(2005, 2009) further elaborated the study of the muscu-
lature morphology of the male genitalia of the Gelechi-
idae, and integrated other morphological characters from
the adult. She divided the family into five subfamilies:
Physoptilinae, Anomologinae, Gelechiinae, Anacampsinae and
the most ‘derived’, Dichomeridinae. Sinev (1992), appar-
ently at least partly independently, suggested nine subfami-
lies in Gelechiidae: Gelechiinae, Anacampsinae, Aristoteliinae,
Metzneriinae, Teleiodinae, Stomopteryginae, Anomologinae,
Brachmiinae and Dichomeridinae. However, he did not give
characters to support this classification, nor did he specify
which genera should be included in each of these subfamilies.
The subdivision of Gelechiidae into tribes is still controver-
sial in the absence of a convincing cladistic analysis based on
a worldwide evaluation. According to Karsholt & Riedl (1996)
seven tribes are currently distinguished within the subfamily
Gelechiinae: Apatetrini, Anomologini, Teleiodini, Gelechiini,
Gnorimoschemini, Anacampsini and Chelariini, whereas the
subfamilies Physoptilinae, Pexicopiinae and Dichomeridinae
are without tribal subdivision. All of these suprageneric groups
can generally be identified only by a combination of various
characters, because parallelism, secondary reductions and ple-
siomorphies are frequent (Huemer & Karsholt, 1999).
Lee & Brown (2008) used DNA data combined with mor-
phological characters to propose phylogenetic relationships of
25 genera of the tribe Litini (formerly Teleiodini) in the Holarc-
tic Region and defined it as a monophyletic clade in subfamily
Gelechiinae. Recently, Nazari & Landry (2012) explored DNA
barcodes as a potential tool to identify species and genera in
the tribe Gnorimoschemini in North America. Ponomarenko
(2005), mainly based on Palaearctic taxa, used male geni-
tal musculature and traditional adult morphological charac-
ters to propose 12 tribes in five subfamilies: Physoptilinae;
Anomologinae (Anomologini, Apatetrini, Aristotelini, Pexi-
copiini); Gelechiinae (Gnorimoschemini, Gelechiini, Litini);
Anacampsinae (Anacampsini, Brachmiini); and Dichomerid-
inae (Dichomeridini, Chelariini, Anarsiini).
Whereas the monophyly of the Gelechiidae has not been
questioned by any recent study its relationships to other
families of Gelechioidea have been debated. Hodges (1998)
found a sister group relationship between Gelechiidae and
Cosmopterigidae based on morphological characters of larva,
pupa and adult. Kaila (2004) confirmed the same relationship
based on a considerably more extensive morphological analysis
and more recently in a molecular analysis based on seven
genes (Kaila et al., 2011). Thus, the sister group position
of these two families seems relatively well supported. How
this pair of families is placed within the Gelechioidea
varies between all these analyses so the sister group to
Gelechiidae + Cosmopterigidae remains unresolved.
Although there is a fair consensus among systematists that
Gelechiidae forms a monophyletic clade (now also including
the Physoptilinae), there is obvious disagreement about how
to subdivide this diverse family. Since Meyrick’s (1925)
original higher classification of the Gelechiidae, the family
has been divided into from three (Hodges, 1986) to nine
subfamilies (Sinev, 1992), and treated variously with different
© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348
combinations of names and levels of tribes and subfamilies.
These inconsistencies have been a result of the different authors
focusing on different, usually narrow, sets of morphological
characters.
The aim of our study is to present higher-level phyloge-
netic relationships of the Gelechiidae based on DNA sequence
analysis, and to compare them with the current morpholog-
ically based hypotheses. There is a need for clarification of
the contents and relationships of the different gelechiid taxa,
which should aid students and researchers of all kinds (includ-
ing taxonomists, applied entomologists, ecologists and amateur
lepidopterists) to associate species with higher taxa in this
species-rich family. We are aware that further data, especially
from faunas in Africa, South America and the Oriental Region,
need to be studied before one can expect a stable system within
the Gelechiidae, but we think that our results as presented here
are a useful step towards that goal.
Materials and methods
Taxon sampling and specimen acquisition
Fifty-seven species exemplars of Gelechiidae and five out-
group taxa were selected for DNA analysis on the basis of
availability and their position within classifications of Hodges
(1998), Ponomarenko (2008b, 2009), Lee et al. (2009) and
Karsholt (2011) (Figs 3–20). These authors list a various
number of subfamilies, of which all except Physoptilinae are
included in this study. Ponomarenko (2008a), Lee et al. (2009)
and Karsholt (2011) divide the largest subfamily, Gelechiinae,
into several tribes, all included in this study. Similarly, Pono-
marenko (2008a) splits four subfamilies into several tribes, all
included in our study. Exemplars from almost all subfami-
lies and tribes were selected in order to test the monophyly
of these taxa, including the Gelechiidae (Table 1). According
to the classification by Hodges (1998) the number of genera
within the 4 therein recognized subfamilies is very uneven,
with 480 genera of Gelechiinae, 6 genera of Dichomeridinae,
21 genera of Pexicopiinae and 1 genus of Physoptilinae. Of
these we analysed 43 genera of Gelechiinae (9.0%), 4 gen-
era of Dichomeridinae (66.7%) and 2 genera of Pexicopiinae
(9.5%). Unfortunately, we were not able to obtain fresh mate-
rial of the monogeneric Physoptilinae, which Hodges (1998)
and Ponomarenko (2009) propose as a sister taxon to all other
gelechiid lineages. The large subfamily Gelechiinae has been
divided into several tribes (see above), but the numbers of
genera in each of these tribes are not available. Some of these
tribes are morphologically rather uniform (e.g. Gnorimoschem-
ini, Anacampsini), whereas others are morphologically diverse
and may not represent monophyletic groups. We selected three
species of Dichomeris, because it is the largest genus of
Gelechiidae with nearly 600 species and numerous generic
synonyms (Hodges, 1986; Ponomarenko, 2009). The taxon
sampling included representatives of 49 currently recognized
genera. Twenty of these are represented by their type species
(40.8%). The sampling was somewhat European biased, but on
 Molecular analysis of Gelechiidae 337

Table 1. List of Gelechiidae species examined, differently categorized within higher classification by earlier authors.

Subfamily
Subfamily Tribus Subfamily Tribus (Pono- Tribus (Pono-
Species Family (European) (European) (American) (American) marenko) marenko)

Scythris limbella Scythrididae – – – – – –

Mompha conturbatella Momphidae – – – – – –
Pancalia schwarzella Cosmopterigidae – – – – – –
Macrobathra chrysotoxa Cosmopterigidae – – – – – –
Limnaecia phragmitella Cosmopterigidae – – – – – –
Anacampsis populella Gelechiidae Gelechiinae Anacampsini Gelechiinae Anacampsini Anacampsinae Anacampsini
Aproaerema anthyllidella Gelechiidae Gelechiinae Anacampsini Gelechiinae Anacampsini Anacampsinae Anacampsini
Mesophleps silacella Gelechiidae Gelechiinae Anacampsini – – Anacampsinae Anacampsini
Carpatolechia notatella Gelechiidae Gelechiinae Teleiodini Gelechiinae Litini Gelechiinae Litini
Stenolechia gemmella Gelechiidae Gelechiinae Teleiodini Gelechiinae Litini Gelechiinae Litini
Pseudotelphusa paripunctella Gelechiidae Gelechiinae Teleiodini Gelechiinae Litini Gelechiinae Litini
Exoteleia dodecella Gelechiidae Gelechiinae Teleiodini Gelechiinae Litini Gelechiinae Litini
Altenia perspersella Gelechiidae Gelechiinae Teleiodini Gelechiinae Litini Gelechiinae Litini
Coleotechnites albicostatus Gelechiidae Gelechiinae Teleiodini Gelechiinae Litini Gelechiinae Litini
Gelechia rhombella Gelechiidae Gelechiinae Gelechiini Gelechiinae Gelechiini Gelechiinae Gelechiini
Athrips pruinosella Gelechiidae Gelechiinae Gelechiini Gelechiinae Gelechiini Gelechiinae Gelechiini
Filatima incomptella Gelechiidae Gelechiinae Gelechiini Gelechiinae Gelechiini Gelechiinae Gelechiini
Neofriseria singula Gelechiidae Gelechiinae Gelechiini – – Gelechiinae Gelechiini
Prolita sexpunctella Gelechiidae Gelechiinae Gelechiini Gelechiinae Gelechiini Anacampsinae Anacampsini
Sophronia semicostella Gelechiidae Gelechiinae Gelechiini Gelechiinae Anacampsini Anacampsinae Anacampsini
Mirificarma flavella Gelechiidae Gelechiinae Gelechiini Gelechiinae Gelechiini Gelechiinae Gelechiini
Psoricoptera speciosella Gelechiidae Gelechiinae Gelechiini – – Gelechiinae Gelechiini
Aroga velocella Gelechiidae Gelechiinae Gelechiini Gelechiinae Gelechiini Gelechiinae Gelechiini
Stegasta bosqueella Gelechiidae – – Gelechiinae Gelechiini Anomologinae Aristoteliini
Chrysoesthia drurella Gelechiidae Gelechiinae Anomologini Gelechiinae Anomologini Anomologinae Apatetrini
Metzneria metzneriella Gelechiidae Gelechiinae Anomologini Gelechiinae Anomologini Anomologinae Anomologini
Aristotelia ericinella Gelechiidae Gelechiinae Anomologini Gelechiinae Anomologini Anomologinae Aristoteliini
Bryotropha terrella Gelechiidae Gelechiinae Anomologini Gelechiinae Gelechiini Anomologinae Aristoteliini
Monochroa hornigi Gelechiidae Gelechiinae Anomologini Gelechiinae Anomologini Anomologinae Anomologini
Eulamprotes wilkella Gelechiidae Gelechiinae Anomologini – – Anomologinae Anomologini
Metanarsia modesta Gelechiidae Gelechiinae Anomologini – – Anomologinae Apatetrini
Psamatocrita osseella Gelechiidae Gelechiinae Anomologini – – Anomologinae Aristoteliini
Megacraspedus Gelechiidae Gelechiinae Anomologini Gelechiinae Anomologini Anomologinae Aristoteliini
Isophrictis striatella Gelechiidae Gelechiinae Anomologini Gelechiinae Anomologini Anomologinae Anomologini
Deltophora sella Gelechiidae Gelechiinae Anomologini Gelechiinae Anomologini Anomologinae Anomologini
Xystophora pulveratella Gelechiidae Gelechiinae Anomologini – – Anomologinae Aristoteliini
Hypatima rhomboidella Gelechiidae Gelechiinae Chelariini Gelechiinae Chelariini Dichomeridinae Chelariini
Nothris verbascella Gelechiidae Gelechiinae Chelariini – – Dichomeridinae Chelariini
Neofaculta infernella Gelechiidae Gelechiinae Chelariini Gelechiinae Chelariini Dichomeridinae Chelariini
Anarsia lineatella Gelechiidae Gelechiinae Chelariini Gelechiinae Chelariini Dichomeridinae Anarsiini
Dactylotula kinkerella Gelechiidae Gelechiinae Apatetrini – – Anomologinae Apatetrini
Caryocolum blandellum Gelechiidae Gelechiinae Gnorimoschemini Gelechiinae Gnorimoschemini Gelechiinae Gnorimoschemini
Scrobipalpa obsoletella Gelechiidae Gelechiinae Gnorimoschemini Gelechiinae Gnorimoschemini Gelechiinae Gnorimoschemini
Scrobipalpopsis petasitis Gelechiidae Gelechiinae Gnorimoschemini Gelechiinae Gnorimoschemini Gelechiinae Gnorimoschemini
Dichomeris juniperella Gelechiidae Dichomeridinae – Dichomeridinae – Dichomeridinae Dichomeridini
Kiwaia lithodes Gelechiidae – – – – Gelechiinae Gnorimoschemini
Brachmia blandella Gelechiidae Dichomeridinae – – – Anacampsinae Brachmiini
Dichomeris latipennella Gelechiidae Dichomeridinae – Dichomeridinae – Dichomeridinae Dichomeridini
Helcystogramma lutatella Gelechiidae Dichomeridinae – Dichomeridinae – Dichomeridinae Dichomeridini
Dichomeris ustalella Gelechiidae Dichomeridinae – Dichomeridinae – Dichomeridinae Dichomeridini
Acompsia cinerella Gelechiidae Dichomeridinae – – – Dichomeridinae Dichomeridini
Pexicopia malvella Gelechiidae Pexicopiinae – (Pexicopiinae) – Anomologinae Pexicopiini
Thiotricha subocellea Gelechiidae Pexicopiinae – – – Anomologinae Aristoteliini
Anisoplaca archyrota Gelechiidaea – – – – – –
Isochasta paradesma Gelechiidaeb,c – – – – – –
Macrenches clerica Gelechiidaed – – – – – –
Xerometra mesophracta Gelechiidaee – – – – – –

a Chelariinae in Edwards (1996).

b Unassigned to subfamily.
c Listed in the genus Aristotelia in Edwards (1996).
d Gelechiinae in Edwards (1996).
e Dichomeridinae in Edwards (1996).

© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348

338 O. Karsholt et al.
the other hand, most named suprageneric gelechiid taxa reach
Europe. Representatives from the Nearctic region, Australia
and New Zealand were also included. A bulk of the samples
was from the DNA collection of Marko Mutanen, but sampling
was supplemented by other authors. Most samples were col-
lected as fresh in absolute alcohol but most air-dried specimens
of less than 3 years old provided DNA of high quality as well.
The identities of sampled species were cross-checked using
the ‘Identification Engine’ tool of the BOLD database (Barcode
of Life Data Systems, http://www.barcodinglife.org/views/
login.php). BOLD database consists of data of the COI 5 gene
region (DNA barcode) of almost all species sampled in this
study and proved therefore to be a useful tool in verifying
identifications. Collection data of specimens are shown in
Table S1 and coverage of DNA sampling with GenBank
accession numbers is provided in Table S2.
Molecular techniques
Usually a few legs of adults, but in some cases body parts
of larvae were used for DNA extraction. The remains of the
extracted specimens were preserved to serve as vouchers that
are deposited in the authors’ institutions. Body parts that were
used in DNA extraction were dried and powdered before the
analysis, and DNA was extracted and purified using DNeasy™
extraction kit and DNeasy Tissue kit(250) (both Qiagen), and
QIAquick PCR Purification kit (50) and Qiagen’s DNeasy
Tissue kit (250) (both Beckman Instruments Inc., California,
U.S.A.), all following manufacturer’s instructions.
In order to infer phylogenetic affinities between the selected
gelechiid taxa, eight gene regions were subjected to DNA
sequencing. Primers for some of these regions were devel-
oped by Wahlberg & Wheat (2008), and primers for gene
regions used in our study are provided in the supplementary
material of their article: 1406 bp region of the cytochrome c
oxidase subunit 1 gene (CO1) of the mitochondrial genome
(sequenced in two parts using LCO-HCO and Jerry-Pat primer
pairs), 850 bp region of the nuclear Carbamoylphosphate syn-
thase domain protein (CAD), 1047 bp region of the nuclear
Elongation Factor-1α (EF-1α) (sequenced in two parts using
Starsky-Monica and EF51.9-EFrcM4 primer pairs), 691 bp
region of the nuclear Glyceraldehyde-3-phosphate dehydro-
genase (GAPDH), 710 bp region of the nuclear Isocitrate
dehydrogenase (IDH), 407 bp region of the nuclear Cytoso-
lic malate dehydrogenase (MDH), 603 bp region of the nuclear
Ribosomal protein S5 (RpS5) and 345 bp region of the nuclear
wingless gene were successfully sequenced. The selection of
these gene regions was based primarily on two rationales.
First, almost universally functioning PCR primers for these
protein coding genes have been developed for Lepidoptera
(see Wahlberg & Wheat, 2008), and second, the evolution of
these gene regions have been shown to be of an informative
rate for inferring family-, subfamily- and tribal-level relation-
ships in other lepidopteran groups (Zahiri et al., 2010, 2012;
Heikkilä et al., 2012; Sihvonen et al., 2011). Altogether, these
gene regions account for a total of 6130 bp. DNA amplification
© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348
and sequencing were carried out using standard PCR and
sequencing techniques following the protocols presented in
Wahlberg & Wheat (2008) in most details, but several dif-
ferent Taq polymerases were applied in PCR (all showing
successful amplification) and PCR reactions were partially
carried out in 10-μL reaction volumes. Sequencing was per-
formed with an ABI 3730 capillary sequencer using dye ter-
minator sequencing kits according to the recommendations of
manufacturers.
Phylogenetic analysis
The sequenced DNA regions represent protein coding genes
whose lengths do not vary between the studied taxa except in
one inserted codon in RpS5 of Bryotropha, rendering the align-
ment straightforward (see e.g. Mutanen et al., 2010; Zahiri
et al., 2010, 2012; Sihvonen et al., 2011). The chromatograms
were checked and sequences aligned manually by eye using
BioEdit v7.0.9.0 (Hall, 1999). However, a fraction of 57 bp
(sites 104–161) was removed in the wingless gene because
of a lot of variation in that area and consequent difficulties
in alignment across sequences in an unambiguous way (the
omitted base pairs are not included in the count of total base
pairs). All sequences were carefully checked for possible cross-
contaminations and other doubtful patterns by constructing a
maximum Likelihood analyses separately for each gene. The
sequence alignments are available in the Table S3 and Gen-
Bank (for accession numbers, see Table S2).
We used five nongelechiid Gelechioidea species as out-
groups. These included three members of Cosmopterigidae
(Limnaecia, Pancalia and Macrobathra), the putative sister
group of Gelechiidae (Hodges, 1998; Kaila, 2004; Kaila et al.,
2011), one genus of Momphidae (Mompha), and one genus
of Scythrididae (Scythris). Scythris was used as a techni-
cal outgroup on which all trees were rooted. The other out-
groups were allowed to freely associate among other taxa,
in order to test the monophyly of Gelechiidae. Various out-
group combinations were used to explore their effects on the
rooting of Gelechiidae. We also explored the influence of
third codon positions by omitting them. Although it had little
effect on the tree topology, the exclusion of third codon posi-
tions resulted in lower supports for many branches. We there-
fore included all data in the eventual analyses. The datasets
were created using the VoSeq program (Peña & Malm, 2012).
The same program was used to create GenBank submission
files.
Phylogenetic analyses were conducted with model-based
Maximum Likelihood (ML) and Bayesian inference meth-
ods. Maximum Likelihood analyses were implemented using
RAxML v7.2.8 Black Box (Stamatakis et al., 2008) at the Phy-
lobench Web portal. These analyses were carried out under the
GTR +  model and the data were partitioned by genes. Sup-
ports for nodes were evaluated with 100 bootstrap replicates
of the data. The analysis was repeated five times to exam-
ine whether rapid algorithms applied in RAxML consistently
found the same global optimum.

Bayesian analyses were carried out in MrBayes v3.1
(Ronquist & Huelsenbeck, 2003) at the CIPRES Web portal
(Miller et al., 2010). Two independent runs of 10 million
generations with three heated and one cold chain and every
1000th tree sampled were performed. Missing nucleotides were
coded as ‘?’. The data were partitioned by gene regions (ten
in total, two genes sequenced in two parts were partitioned
by these regions). The tree priors were set to ‘variable’.
The branch lengths were allowed to vary under a relaxed
clock model with an uncorrelated lognormal distribution.
A nucleotide substitution model GTR +  was applied for
each partition using the ‘unlink’ command in MrBayes.
Node supports were estimated by posterior probabilities. The
value of 0.02 of standard deviation of split frequencies was
reached at about 1.8 million generations and at 10 million
generations the value was about 0.013, implying that the
two runs were almost perfectly converged at this point and
the run was stopped. Convergence was determined when
the standard deviation of split frequencies went below 0.05
and the PSRF (Potential Scale Reduction Factor) approached
1, and both runs had properly converged to a stationary
distribution after the burn-in stage (which was 2000 sampled
generations).
Results
Monophyly of Gelechiidae, its subfamilies and tribes
The five repeats of ML analyses produced trees with the
same topologies, only with some variation in the bootstrap
support values (hereafter abbreviated BS). For some basally
arising nodes BS was less than 50%. Figure 1 shows the
ML tree with the observed range of node support values.
Nodes that were not supported in all the repeats are indicated
as ‘no’ in the node support labels. The Bayesian analysis
yielded a well-resolved tree nearly identical to that of ML
in topology (Fig. 2), but usually with significantly higher
posterior probabilities (hereafter abbreviated PP). We interpret
PP values conservatively and nodes supported when they reach
the value of 0.8 or more. The only difference in the topology
between the ML and Bayesian tree is in the level of resolution
within Anomologinae (Figs 1, 2).
Both ML and Bayesian analyses found monophyly of
Gelechiidae very strongly supported and the sister group rela-
tionship of Cosmopterigidae + Gelechiidae well-supported as
well, although our present nongelechiid taxon sampling was
too limited to reliably estimate the sister group relationship
between Gelechiidae and Cosmopterigidae. However, Cos-
mopterigidae have consistently been found as a sister group of
Gelechiidae in several recent analyses of phylogenetic relation-
ships of Gelechioidea (Hodges, 1998; Kaila, 2004; Mutanen
et al., 2010; Kaila et al., 2011).
Of the three included gelechiid subfamilies, Gelechiinae,
Pexicopiinae and Dichomeridinae sensu Hodges (1998) and
Karsholt (2011) (i.e. including Anacampsini and Chelariini),
only Dichomeridinae were found to be monophyletic, with the
© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348
Molecular analysis of Gelechiidae 339
tribe Brachmiini nested deeply within it. Both Dichomeridinae
and Pexicopiinae were found to be nested within the Gelechi-
inae, rendering that subfamily paraphyletic. The BS values
for some of the most basally arising nodes of the broadly
delineated Gelechiinae are low. However, Bayesian analysis
gives a high PP (0.95) for the clade comprising all gelechiid
taxa except Anacampsini and Chelariini. Of the five sub-
families included by Ponomarenko (2008a, 2009) Physoptili-
nae were not included into this study, and of the remaining
four subfamilies only Gelechiinae sensu Ponomarenko (2008a,
2009) and Dichomeridinae were found to be monophyletic.
Both Anacampsinae and Anomologinae as delimited by Pono-
marenko, were split into three distinct clades rendering these
subfamilies polyphyletic.
A weakly supported (32–41% BS, 0.72 PP) clade com-
prising Anacampsini + Chelariini (sensu Karsholt, 2011)
(Gelechiinae) forms the sister clade for the remaining gelechiid
taxa in all analyses. The next branching event is between
the clade of Dichomeridinae and Pexicopiinae + Gelechiinae
(sensu Karsholt, 2011 except for the above-mentioned exclu-
sion of Chelariini and Anacampsini). This clade receives a low
BS (26–33%), but relatively high PP (0.95). Within this clade,
the Dichomeridinae (sensu Karsholt, 2011) is very strongly
supported as a monophyletic group (96–99% BS, 1.0 PP). The
next lineage comprises Pexicopiinae, and Apatetrini (Gelechi-
inae) (sensu Karsholt, 2011) plus two taxa (Chrysoesthia and
Metanarsia) placed in Anomologini in Karsholt (2011). Aniso-
placa, tentatively placed in Chelariini by Edwards (1996), is
a close relative of Pexicopia, the type genus of Pexicopiinae
(100% BS and 1.0 PP). The sister of that clade (with 38–61%
BS and 1.0 PP) comprises of Metanarsia, Chrysoesthia and
Dactylotula, which is a monophylum equal to Apatetrini sensu
Ponomarenko (2005, 2009), strongly supported with 100%
BS, 1.0 PP. The next clade with Thiotricha + Macrenches
and most Gelechiinae excluding Chelariini, Anacampsini,
Apatetrini and a few Anomologini (sensu Karsholt, 2011) is
well supported (79–91% BS, 1.0 PP). An affinity between
Thiotricha and Macrenches is a novel group which receive
some BS support (20–59%), and high PP support (0.96). The
core Gelechiinae (sensu Karsholt, 2011) without the above-
mentioned taxa receives low BS support (18–50%), but high
PP support (0.95). This clade is divided into Anomologini
(sensu Karsholt, 2011) including Bryotropha, and Gelechiini +
Gnorimoschemini + Litini (with Xystophora of Anomologini
placed here as well).
Tribal division of Gelechiinae by Karsholt (2011) is gen-
erally well supported. Anacampsini, Chelariini, Anomologini,
Gnorimoschemini and Litini are all well supported as mono-
phyletic groups, respectively (with the exceptions mentioned
above), but Gelechiini appears paraphyletic because both Gno-
rimoschemini and Litini are nested within it. Aristotelini
(Gelechiinae), as recognized by Ponomarenko (2008a, 2009),
is polyphyletic in our cladograms, with members scattered
in Anomologini and Gelechiini sensu Karsholt (2011). Pono-
marenko (2008a, 2009) also divides Anacampsinae into two
tribes, Anacampsini and Brachmini, of which the only included
340 O. Karsholt et al.

Fig. 1. Maximum Likelihood Tree for Gelechiidae, implemented under GTR +  model and rooted on Scythris (Gelechioidea, Scythrididae).
Numbers in nodes indicate the observed range of support values, and numbers with ‘no’ indicate nodes not supported in all the repeats.

© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348

 Molecular analysis of Gelechiidae 341

Fig. 2. Bayesian Tree, rooted on Scythris (Gelechioidea, Scythrididae), for Gelechiidae. The tree is based on two independent runs of 10 million
generations implemented under GTR +  model. Numbers in nodes indicate Posterior Probability values (PP).

member of Brachmini is nested within Dichomeridini in our morphological evidence on which they have been based, and
results. suggest a revised classification.

Discussion
In the first section of discussion we compare our molecular
results with previous hypotheses of phylogeny and suggested
Anacampsinae
The first split within the Gelechiidae is between the seven
taxa from Aproaerema to Anarsia and the remaining 45 taxa
studied (Figs 1, 2). This clade includes the tribes Anacampsini

© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348

342 O. Karsholt et al.

Figs 3–20. Adult representatives of Cosmopterigidae and main Gelechiidae lineages observed in the study. 3, Pancalia schwarzella; 4, Anacampsis
populella; 5, Hypatima rhomboidella; 6, Dichomeris juniperella; 7, Helcystogramma lutatella; 8, Chrysoesthia drurella; 9, Pexicopia malvella;
10, Thiotricha subocellea; 11, Bryotropha terrella; 12, Aristotelia ericinella; 13, Eulamprotes wilkella; 14, Athrips pruinosella; 15, Sophronia
semicostella; 16, Filatima incomptella; 17, Scrobipalpa obsoletella; 18, Gelechia rhombella; 19, Carpatolechia notatella; 20, Stenolechia gemmella.

and Chelariini of Lee et al. (2009) and Karsholt (2011), Dichomeridinae. Ponomarenko also separated Anarsia and
which we consider here to be sister groups within the sub- related genera into the tribe Anarsiini of Dichomeridinae
family Anacampsinae. This is in opposition to Ponomarenko (Ponomarenko 2008a, 2009). We found no support for this
(2008a, 2009) who treats Chelariini as belonging to the separation as Anarsia is inside the Chelariini in our analysis.

© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348


Dichomeridinae
The clade consisting of Dichomeris, Acompsia, Brachmia
and Helcystogramma in our study receives high support and
corresponds to Dichomeridinae of Hodges (1986), Lee et al.
(2009), and Karsholt (2011). This result is quite different from
Ponomarenko (2005, 2008a, 2009) who included Chelariini
in Dichomeridinae and placed Brachmia in a tribe of its
own, Brachmiini, in Anacampsinae. This conclusion was based
mainly upon muscle protractors of phallus m5 forming a
single bundle, whereas it is divided into two bundles in
Dichomeridinae. This echoes Edwards (1996) who recognized
a subfamily Brachmiinae. That group was, however, obviously
heterogeneous as one of its two constituent genera, Anatiplora,
has been suggested to be associated with Autostichidae (Kaila,
2004; Kaila et al., 2011). We found Brachmia nested deeply
within the Dichomeridinae, next to Helcystogramma. This
placement also seems to be well supported by morphology.
Brachmia, Helcystogramma and Acompsia (Dichomeridini)
share the following characters: the shape of the forewing is
sub-triangular; the cellular cross-vein on hindwing is absent;
male genitalia have a free, strongly inflated phallus; female
genitalia possess a large ostium bursae that is submerged to
segment VII.
Hodges (1986) based his concept of the Dichomeridinae
mainly on the structures of the abdominal support system
of sternum II listing the following traits: apodemes reduced;
lateral process at base of apodemes absent; sternal rods
comparatively short; anterior margin of sternum II between
apodemes concave and sclerotised. However, as pointed out to
us by (K. Gregersen, personal communication) the abdominal
support system in Anacampsini (but not in Chelariini) is
very similar to that of Dichomeridinae, and there is also
overall similarity in the male genitalia between these two
groups. However, according to our results, a clade consisting
of Anacampsini and Dichomeridinae but without Chelariini
would be paraphyletic. Kaila (2004) discussed the validity
of phylogenetic signal of these structures of the thoraco-
abdominal support system both within Gelechioidea, and
Lepidoptera in general, and concluded that they are quite
plastic and thus should not be over-emphasized in inferring
interrelationships of taxa. Our present finding seems to
support this. The overall similarity of the genitalia between
Dichomeridinae and Anacampsini may be interpreted as
symplesiomorphic, with Chelariini displaying apomorphic
features.
Anomologinae
The recovery of a lineage consisting of Pexicopia and
Anisoplaca, with Dactylotula, Chrysoesthia and Metanarsia
as its sister group is at first glance an odd assemblage,
but not that surprising. Even though Anisoplaca has been
placed in the Chelariini, it appears to be a close relative
of Pexicopia in the Pexicopiinae also by its morphology
(R. Hoare, personal communication). Ponomarenko (2008a,
© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348
Molecular analysis of Gelechiidae 343
2009) included Pexicopiinae as a tribe in Anomologinae,
in which she also included Anomologini, Apatetrini and
Aristoteliini. She also suggested that Apatetrini included
Chrysoesthia and Metanarsia, which had been included in
Anomologini by Karsholt (2011) and others. Pexicopiini and
Apatetrini are sister groups according to our results. We
consider them as tribes of one subfamily, Apatetrinae, as this
subfamily possesses at least one synapomorphy: the antenna
with a fully developed pecten. Although a pecten is generally
a homoplastic character among Gelechioidea, its presence is
nearly always consistent with relationships at the subfamily
level (Kaila, 2004). Within Gelechiidae a fully developed
antennal pecten is only present in Apatetrinae (as defined here).
When present outside of the Apatetrinae it consists of only a
single bristle. In our view, the presence of a fully developed
antennal pecten supports the suggestion that Pexicopiini should
be considered subordinate to Apatetrinae. This classification
is also in accordance with other recent ones where fewer
subfamilies have been favoured over additional splittings. With
the exception of Anarsiini, this view is in line also with
Ponomarenko (2009).
Thiotrichinae subfam.nov.
Type genus: Thiotricha
We found that Thiotricha and Macrenches formed a
monophyletic lineage, not subordinate to other lineages.
Macrenches is listed by Edwards (1996) in Gelechiinae.
Based on our observation, M. clerica (Rosenstock) also
shares morphological similarities with Thiotricha (e.g. a
large, posteriorly bifurcate sternum VIII, a small tergum
VIII, a sickle-shaped signum, and the male with rather long
ciliated antennae) although obviously these are not congeneric
(e.g. the finger-formed anellus projections characteristic to
Thiotricha are missing in Macrenches). Moreover, the larva of
M. clerica is a web-spinner feeding on Mimosaceae (Common,
1990), whereas larvae of Thiotricha as far as known feed
from within a portable case on different plant families
such as Symplocaceae (e.g. Ueda & Fujiwara, 2005). The
pupa of Thiotricha is characterized by the following traits
that are unusual in Gelechiidae: the head is armed with
small spines; the mesothorax has conical lateral corners to
varying extent; antennae approach each other but do not
meet mesially; abdominal segments 7 and 8 have posterior
rows of minute teeth (Ueda & Fujiwara, 2005, L. Kaila,
personal observation). In addition to Thiotricha we assign the
following genera to Thiotrichinae: Macrenches Meyrick, 1904,
Palumbina Rondani, 1876 (= Thyrsostoma Meyrick, 1907) and
Polyhymno Chambers, 1874.
Thiotrichine moths are overall slender-winged Gelechiidae,
with the forewings often with longitudinal streaks and apically
with distinctive markings; the apex is often upturned. They are
morphologically diverse, even within the type genus Thiotricha
(Ueda & Fujiwara, 2005), making it difficult to characterize
them. Thiotrichinae are distributed primarily in warmer areas
of both the old and the new world, and are most diverse in
Asia. The group is in much need of revision. Relevant literature
344 O. Karsholt et al.
on the group include, beside of the catalogues by Meyrick
(1925) and Gaede (1937), Janse (1949–1963) (South Africa),
Ivinskis et al. (1984), Huemer (1993) and Ueda & Fujiwara
(2005) (Palaearctis) and Sattler (1982) (genus Palumbina).
Thiotricha was synonymized with Polyhymno Chambers by
Ponomarenko (2009) without further comments. We could
not include Polyhymno in our molecular analysis due to
lack of fresh material, but we compared the type species
P. longistrigella Chambers morphologically with species of
Thiotricha. Thiotricha share with Polyhymno a well-developed
pterostigma in the forewing, a large sternum VIII and a small
tergum VIII. However, sternum VIII is not posteriorly bifurcate
in Polyhymno, which has moreover a simple valve with
reduced anellus projections. Polyhymno also has shorter ciliae
at the male antenna. Contrary to Thiotricha (whose known
larvae feed from within a case, e.g. Ueda & Fujiwara, 2005)
larvae of Polyhymno are leaf-spinners and leaf-webbers in
Fabaceae (Busck, 1900). These morphological differences and
the different feeding habits among these two genera support our
opinion that Thiotricha needs to be resurrected. We herewith
remove Thiotricha from synonymy with Polyhymno and treat
it as a separate genus (rev. status). Had the two genera been
synonyms, we would have had to base the new subfamily name
on Polyhymno which is the oldest of them (ICZN, 1999 article
11.7.1).
We prefer to base it on Thiotricha, as this genus was included
in our molecular study, and with nearly 100 described species
it is the largest and most well-known of the genera here
assigned to Thiotrichinae. Even though the statistical support
for uniting Thiotricha and Macrenches is not very high, we
assume that this may be due to the scarce taxon sampling in this
species-rich group where these two exemplar taxa are about as
different from each other as possible. Addition of taxa would
more likely strengthen than weaken this association (L. Kaila,
personal observation). The position of this lineage, together
with the morphological evidence therefore supports that this
clade should be treated as a subfamily of its own.
In search for available family-group names for what we
now term Thiotrichinae we considered the name Palumbininae
Chapman, 1902, which was included in the Gelechiidae by
Nye & Fletcher (1991). Its type genus Palumbina Rondani,
1876 was not included in the present study, but it is
morphologically similar to Thiotrichinae. Ponomarenko (2005,
2009) rejected Palumbininae as a nomen nudum. It was
apparently not the intention of Chapman (1902) to propose a
new subfamily, and the action was probably due to a printing
error, which is corrected by pencil in the copy of volume
of The Entomologist in question held by the library of the
Entomological Department of the Natural History Museum in
London (K. Sattler, personal communication). We therefore
support Ponomarenko’s (2005, 2009) opinion and follow it
here.
Diagnosis. Within Gelechiidae, Thiotrichinae is defined by
the molecular differences shown in this study as well as
morphological characters such as a well-developed pterostigma
© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348
in the forewing (also present in, e.g., some Litini), a large
sternum VIII and a reduced tergum VIII, signum a plate (e.g.
sickle-shaped, jar-shaped or occasionally missing), and the
male antennae with rather long cilia (shorter in Polyhymno).
Thiotrichinae subfam.nov. has been regisetered in ZooBank.
Anomologinae
A clade containing the taxa from Bryotropha to Eulamprotes
corresponds to Anomologini in Gelechiinae s.l. of Karsholt
(2011), except for a few genera that are now associated
with other subfamilies, e.g. Chrysoesthia and Metanarsia in
Apatetrinae and Xystophora in Gelechiinae s.s. This clade
is the sister group of Gelechiinae s.s., and we consequently
propose to treat it equally as Anomologinae s.s. Ponomarenko
(2008a, 2009) suggested a subfamily Anomologinae, but in a
wider sense, including also Apatetrini and Pexicopiini which
according to our findings, with good support, are not members
of this group. Anomologinae s.s. as proposed here is a rather
compact clade in our analysis. We find no support to divide it in
Anomologini and Aristoteliini as suggested by Ponomarenko
(2008a, 2009). Unfortunately we were unable to include
Anomologa Meyrick, 1926, the type genus of Anomologinae,
in our study. In both the ML and Bayesian analyses Bryotropha
came out as a sister group to the remaining Anomologinae s.s.,
although only in the latter with high support. This appears
reasonable as Bryotropha differs in several aspects from
other Anomologinae, and its closest relatives remain unknown
(Rutten & Karsholt, 2004; Karsholt & Rutten, 2005).
Xerometra mesophracta (Turner) from Tasmania, which was
listed in the Dichomeridinae by Edwards (1996), is here
nested in Anomologini. It was later removed from Xerometra
and left unplaced in the Gelechiidae (Li & Sattler, 2012;
Su et al., 2012), although it was supposed to belong in
the Anomologinae (T. Edwards, personal communication).
According to its morphology we consider it to be a close
relative of Megacraspedus.
We consider the recognition of Anomologinae as a subfamily
to be a more conservative classificatory act than any other
way that could be derived from our well supported result.
Recognition of it is also in accordance with Ponomarenko
(2009).
Gelechiinae
Our data support the monophyly of a clade containing
the studied taxa from Xystophora to Exoteleia (Fig. 1),
namely 21 out of 52 studied Gelechiidae taxa. This clade
corresponds to the definition of the subfamily Gelechiinae
sensu Ponomarenko (2008b, 2009), apart from Ponomarenko’s
inclusion of Xystophora in Anomologinae and Sophronia
in Anacampsinae. Their placement here seems not to be
contradicted by their morphology either, which supports
their previous classification as Gelechiinae by Hodges (1986,
1998). The Gelechiinae in this sense (s.s.) are characterized

by one synapomorphy: tergum VIII and sternum VIII in
the male have free ‘flaps’ that are formed of an outer
sclerotized part belonging to tergum VIII and sternum VIII,
and an inner unsclerotized membrane, which is formed by
the intersegmental membrane between segment VIII and IX,
so they are not united to a ring (Huemer & Sattler, 1995,
N.P. Kristensen, personal communication). Tergum VIII is in
many species in this clade tongue-shaped, and sternum VIII
is usually broad, sometimes with paired anterior apophyses,
namely rod-like lobes (Hodges, 1998; Huemer & Karsholt,
1999). During copulation tergum VIII and sternum VIII can
open widely, allowing the male to come closer with its genitalia
to the female (Huemer & Sattler, 1995). The valvae (cucullus
and sacculus) in these genera show a tendency to becoming
shorter and more slender (especially in the Litini), and it is
likely that these free flaps compensate for this lost ability to
grip the female (O. Karsholt, unpublished data).
The condition with separate tergum and sternum VIII is pre-
vailing in Lepidoptera, and their fusion as a ring, as in figs
53–58 in Huemer & Sattler (1995), is considered a synapo-
morphy for the Gelechiidae by these authors. The condition
found in Gelechiinae as understood in the present paper is
thus a further modification. The inclusion of Xystophora into
Gelechiinae s.s. is new. This genus has hitherto been placed in
the Anomologinae (-ini), without any close relative. Presence
of such a complex structure further supports the molecular-
based finding of the monophyly of this clade.
Sophronia has hitherto been associated with various taxa
within Gelechiidae. Karsholt & Riedl (1996) placed it in the
Dichomeridinae. Corley (2001) suggested that only some of the
species in the genus belong to that subfamily, and he erected a
new genus Pseudosophronia for these, whereas the remaining
Sophronia species were placed in the Gelechiinae. Otherwise,
Ponomarenko (2008b) and Lee et al. (2009) retained Sophronia
in Anacampsini. Our study suggests a well-supported sister
group relation with Mirificarma, which has not been suggested
before.
The current competing tribal classifications of Gelechiinae
s.s. agree in separating the subfamily into three tribes: Litini
(= Teleiodini), Gelechiini and Gnorimoschemini (e.g. Huemer
& Karsholt, 1999, 2010; Ponomarenko, 2008a, 2009; Lee
et al., 2009). However, this is unsupported by our results,
as Gelechiinae (-ini) is paraphyletic with respect to Litini
and Gnorimoschemini. But, as our data are insufficient in
proposing a new and better classification, we suggest, for
the time being, the continuing use of the established tribes.
We further stress the need of a more detailed study with a
considerably increased taxon sampling, to be able to suggest a
satisfying tribal subdivision for Gelechiinae s.s.
Physoptilinae
We could not include Physoptilinae in this study due to the
lack of fresh material. This small subfamily includes only the
type genus, Physoptila Meyrick, 1914 with seven species from
the Indo-Australian region. It was defined by two characters,
© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348
Molecular analysis of Gelechiidae 345
considered as ‘parallelisms’ in the context of the superfamily
Gelechioidea by Hodges (1998): an ankyloid phallus and
rows of spines on the pupal tergites. Additionally a hyaline
area on the hindwing was mentioned by Hodges (1998) as a
possible autapomorphy [such a hyaline area is also found in the
hindwing of several species of Anarsia (O. Karsholt, personal
observation)]. The abdominal support system of Physoptila
has sternum II with paired venulae and apodemes, as in
most Gelechiidae. Although characters of the larva have not
been described, larvae as far as known are stem-borers on
Lecythidaceae (Meyrick, 1914). The phylogenetic relationships
of this subfamily remain unclear, pending further study.
Patterns of evolution of life history traits
In this section we discuss how the life-history traits of
Gelechiidae are correlated with the phylogenetic hypothesis
suggested here.
Gelechiidae is one of the most diverse lepidopteran families
in terms of species number. Much of this success can
probably be attributed to the exceptional diversity of life-
history strategies within the family, in combination with
frequent mono- or oligophagous species. Many species are
external feeders at the larval stage, but the larvae stay in a
shelter of some kind, often between leaves that have been
tied together, or within leaf rolls. Larvae of certain groups,
especially in Gnorimoschemini, are known to rest in silken
tubes underground from where they emerge to consume leaves
of the host plants (Kaitila, 1996; Huemer & Karsholt, 2010).
Others are internal feeders to varying degree: some bore into
flowerheads of host plants, others in stems or roots. Some
species of Caryocolum and Gnorimoschema (Gelechiinae:
Gnorimoschemini) induce galls to their host plant. Leaf-mining
is a specialized trait that appears to have independently evolved
multiple times within Gelechiidae. This is an intriguing finding
as this mode of life appears rare in Lepidoptera general,
although when acquired it has often led to very species-rich
radiations (Kaila et al., 2011). The portable cases of the larvae
in the genus Thiotricha (Thiotrichinae) constitute a unique trait
within Gelechiidae. According to Heikkilä & Kaila (2010) and
Kaila et al. (2011) this feature is known only in four other
occasions within Gelechioidea.
According to our results, little of this diversity in larval
mode of life has direct phylogenetic correlation at the higher
(subfamily) level within Gelechiidae. Even closely related
groups can display quite different larval behaviour. For
instance, the larvae of Metzneria (Anomologinae) live in
flowerheads of Asteraceae, whereas larvae of the closely
related genus Monochroa are leaf or stem miners. Monochroa
species are apparently mono- or oligophagous, but at the
generic level the array of host plants encompasses many
distantly related plant families, including monocotyledons,
like Carex spp. which are the host plant of M. ferrea
(Frey), and several distantly related dicotyledon plant families
(Primulaceae, Polygonaceae, Rosaceae etc.) and even ferns
which are the host plant of M. cytisella (Curtis). It appears
346 O. Karsholt et al.
that, in general, any phylogenetic signal related to larval mode
of life or host plant families is at a tribal, generic or lower
level. An example of a feature appearing at the tribal level is
the host selection of Litini larvae that have a general tendency
to feed on trees, both broadleaved and coniferous.
Given the obvious lack of correlation between particular life-
history traits and phylogeny in Gelechiidae, it is not surprising
that no apparent ‘pest clade’ can be identified. Each of the
most notorious pests has its own mode of life, and the pest
species are scattered among subfamilies. The pink bollworm
(Pectinophora gossypiella; Apatetrinae: Pexicopiini) bores into
flowerheads of its host plants in Malvaceae. Angumois grain
moth (Sitotroga cerealella; Apatetrinae: Pexicopiini) bores into
seeds of its poacean hosts and is a serious pest of stored cereals.
The potato tuber worm Phthorimaea operculella (Gelechiinae:
Gnorimoschemini) bores in the roots and tubers of potato and
tobacco. The tomato leaf miner Tuta absoluta (Gelechiinae:
Gnorimoschemini) mines leaves and cambium of stems of
tomato and other Solanaceae. Other pest species included in
this study are Recurvaria nanella (Gelechiinae: Litini) the larva
of which is first a leaf miner, later living in spun shoots,
leaves and flower buds of Rosaceae, and Anarsia lineatella
(Anacampsinae: Chelariini) which is a borer in shoots, twigs
and fruits of Rosaceae. Among the numerous gelechiid pests
listed by Zhang (1994) the most serious ones are miners and
borers, but these strategies are found scattered throughout all
subfamilies of Gelechiidae.
Conclusions
Based on our findings, and considering them against earlier
classifications to make a minimum number of changes, we sep-
arate the family Gelechiidae into seven subfamilies (Physoptili-
nae, Anacampsinae, Dichomeridinae, Apatetrinae, Thiotrichi-
nae, Anomologinae, Gelechiinae) (Fig. 2). The Pexicopiinae is
according to our data a subordinate clade (Pexicopiini) within
the Apatetrinae.
For the clade consisting of Thiotricha and Macrenches
we propose Thiotrichinae subfam.nov. based on the genus
Thiotricha Meyrick, originally described from New Zealand
(Meyrick, 1886). Our findings moreover have the consequences
that Brachmini should again be considered a synonym of
Dichomeridinae, and Anarsiini should again be considered a
synonym of Chelariini.
We find that life-history patterns are not generally correlated
with subfamilies. Such correlations are, though, often evident
at lower systematic levels – tribal level and lower. Gelechiid
pest species do not belong to a single clade but are scattered
among different lineages.
Supporting Information
Additional Supporting Information may be found in the online
version of this article under the DOI reference:
10.1111/syen.12006
© 2013 The Royal Entomological Society, Systematic Entomology, 38, 334–348
Table S1. Collection data of sequenced specimens.
Table S2. Specimen list with gene regions. Numbers
indicate the length of a DNA molecule, i.e. base pairs.
GenBank accession numbers are indicated in parenthesis.
Lack of success in DNA sequencing is indicated by
‘missing’.
Table S3. Sequence alignments used in the study. For
wingless, both the used alignment with a region of 57 bp
removed (named wgl_nogap) and the complete sequence
are provided.
Acknowledgements
We are grateful to Keld Gregersen, Sorø, Denmark for
fruitful discussions on gelechiid systematics, and to Ted
Edwards, Australian National Insect Collection, Canberra,
Australia, Niels Peder Kristensen and Thomas Pape, Zoological
Museum, Natural History Museum, Copenhagen, Denmark,
Jean-Francois Landry, Agriculture and Agri-Food Canada,
Ottawa, Canada and Klaus Sattler, The Natural History
Museum, London, U.K., for providing us with valuable
information. Maria Heikkilä, Finnish Museum of Natural
History, Finland, kindly helped with Bayesian analyses and
Jeremy deWaard, Biodiversity Institute of Ontario, Guelph,
Canada, provided us a DNA barcode sequence of Physoptila.
The study was financially supported by the Academy of
Finland, grant #1110906 to Lauri Kaila. Laura Törmälä
is cordially thanked for the help in lab work. We thank
Thomas Simonsen, The Natural History Museum, London,
U.K., Richard L. Brown, Mississippi Entomological Museum,
Mississippi, USA and two anonymous referees for helpful
comments on the manuscript.
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