A new molecular phylogeny offers hope for a stable family level classification of the Noctuoidea (Lepidoptera)

Abstract

Zahiri, R., Kitching, I. J., Lafontaine, J. D., Mutanen, M., Kaila, L., Holloway, J. D. & Wahlberg, N. (2010). A new molecular phylogeny offers hope for a stable family level classification of the Noctuoidea (Lepidoptera). —Zoologica Scripta, 40, 158–173.
To examine the higher level phylogeny and evolutionary affinities of the megadiverse superfamily Noctuoidea, an extensive molecular systematic study was undertaken with special emphasis on Noctuidae, the most controversial group in Noctuoidea and arguably the entire Lepidoptera. DNA sequence data for one mitochondrial gene (cytochrome 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) were analysed for 152 taxa of principally type genera/species for family group taxa. Data matrices (6407 bp total) were analysed by parsimony with equal weighting and model-based evolutionary methods (maximum likelihood), which revealed a new high-level phylogenetic hypothesis comprising six major, well-supported lineages that we here interpret as families: Oenosandridae, Notodontidae, Erebidae, Nolidae, Euteliidae and Noctuidae.

Full text

A new molecular phylogeny offers hope for a stable family
level classification of the Noctuoidea (Lepidoptera)
REZA ZAHIRI,IAN J. KITCHING,J.DONALD LAFONTAINE,MARKO MUTANEN,LAURI KAILA,JEREMY D.
HOLLOWAY &NIKLAS WAHLBERG
Submitted: 5 May 2010
Accepted: 30 September 2010
doi:10.1111/j.1463-6409.2010.00459.x
Zahiri, R., Kitching, I. J., Lafontaine, J. D., Mutanen, M., Kaila, L., Holloway, J. D. &
Wahlberg, N. (2010). A new molecular phylogeny offers hope for a stable family level
classification of the Noctuoidea (Lepidoptera). — Zoologica Scripta,00, 000–000.
To examine the higher level phylogeny and evolutionary affinities of the megadiverse
superfamily Noctuoidea, an extensive molecular systematic study was undertaken with spe-
cial emphasis on Noctuidae, the most controversial group in Noctuoidea and arguably the
entire Lepidoptera. DNA sequence data for one mitochondrial gene (cytochrome oxidase
subunit I) and seven nuclear genes (Elongation Factor-1a,wingless, Ribosomal protein S5,
Isocitrate dehydrogenase, Cytosolic malate dehydrogenase, Glyceraldehyde-3-phosphate
dehydrogenase and Carbamoylphosphate synthase domain protein) were analysed for 152
taxa of principally type genera ⁄species for family group taxa. Data matrices (6407 bp total)
were analysed by parsimony with equal weighting and model-based evolutionary methods
(maximum likelihood), which revealed a new high-level phylogenetic hypothesis compris-
ing six major, well-supported lineages that we here interpret as families: Oenosandridae,
Notodontidae, Erebidae, Nolidae, Euteliidae and Noctuidae.
Corresponding author: Reza Zahiri, Department of Biology, Laboratory of Genetics, University
of Turku, 20014 Turku, Finland. E-mail: reza.zahiri@utu.fi
Ian J. Kitching and Jeremy D. Holloway, Department of Entomology, Natural History Museum,
Cromwell Road, London SW7 5BD, UK. E-mails: i.kitching@nhm.ac.uk; j.holloway@nhm.ac.uk
J. Donald Lafontaine, Canadian National Collection of Insects, Arachnids and Nematodes, Agricul-
ture and Agri-Food Canada, K. W. Neatby Building, Central Experimental Farm, Ottawa,
Ontario, Canada K1A 0C6. E-mail: lafontained@agr.gc.ca
Marko Mutanen, Department of Biology, University of Oulu, P.O. Box 3000, FIN-90014 Oulu,
Finland. E-mail: marko.mutanen@oulu.fi
Lauri Kaila, Zoological Museum, Finnish Museum of Natural History, P.O. Box 17, FI-00014
University of Helsinki, Helsinki, Finland. E-mail: lkaila@mappi.helsinki.fi
Niklas Wahlberg, Department of Biology, Laboratory of Genetics, University of Turku, 20014
Turku, Finland. E-mail: niklas.wahlberg@utu.fi
Introduction
The classification of the diversity of life on Earth is one of
the major ongoing undertakings of human society (Wilson
2000), but is still far from completion, both in terms of
the inventory of species and of the classification of those
species in a hierarchical system that has a phylogenetic
basis. Although there has been some discussion of aban-
doning the 250-year-old Linnaean system for such classifi-
cation, focusing more on genetic diversity, most
researchers still prefer to use this hierarchical system. It
provides a framework for access to a massive information
resource on the biology, ecology and economic impor-
tance of all species, perhaps epitomized today by the
Encyclopaedia of Life initiative (Wilson 2003). The use of
molecular data, in particular DNA sequences, is becoming
increasingly important for testing and improving classifi-
cations, especially for highly diverse groups of organisms
such as insects.
Lepidoptera are one of the four major Orders of insects,
and Noctuoidea are the largest superfamily within the
Order with between 42 000 (Heppner 1991) and 70 000
(Kitching & Rawlins 1998) described species. The next
most numerous superfamily is Geometroidea with about
21 500 species, followed by Pyraloidea, Papilionoidea and
Gelechioidea each with between 15 000 and 16 000 spe-
cies (Heppner 1991). In contrast, the total number of ter-
restrial vertebrates is approximately 21 500 species
(Maddison 2007).
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 2010 1
Zoologica Scripta

The monophyly of Noctuoidea, based on the presence
of a single apomorphic character, the metathoracic tymp-
anal organ (Miller 1991) and its associated abdominal
structures, seems well established (Kitching & Rawlins
1998; Mitchell
et al.
2006). However, the limits and con-
tent of the constituent families, and the evolutionary rela-
tionships amongst and within these, are very poorly
understood (Mitchell
et al.
2006).
Noctuoid species are placed in approximately 4200 gen-
era (Kitching & Rawlins 1998) but there are numerous
undescribed species, particularly from tropical regions. For
example, the noctuoid species total for Borneo now stands
at about 2800, an increase of over 50% on a total esti-
mated from a previous survey (1980) of the collections of
The Natural History Museum in London (J. D. Holloway,
unpublished data). This increase is the result of a quarter
of a century of taxonomic effort on a major input of fresh
survey material from Borneo, leading to a series of mono-
graphs on the Macrolepidoptera, those on the noctuoids
mostly cited in this paper. Taking as a minimum the cal-
culations of the total Bornean Lepidoptera fauna by Rob-
inson & Tuck (1993, 1996), this figure for noctuoids
represents about one quarter of that total. The proportion
of Noctuoidea in the global total is likely to be similar.
The larvae of many noctuoid genera include army-
worms, cutworms, bollworms and stem borers, that collec-
tively have a massive economic impact annually (Kitching
1984). The adults of other genera damage fruit crops by
piercing the skins to suck juices (Ba
¨nziger 1982). Noctu-
oids constitute one quarter of the approximately 6000
Lepidoptera species noted to be of economic importance
by Zhang (1994). Though many of these can be assigned
to what Mitchell
et al.
(2006) termed the ‘pest clade’, many
more are distributed across the whole superfamily in over
500 genera (Zhang 1994). Therefore, resolution of stable,
extrapolative higher level classificatory structure for the
superfamily may prove to be an important prerequisite for
studies of pest bionomics across the group.
Numerous classifications of the family groups of Noctu-
oidea have been proposed. The fundamental distinction
between the different systems is based on the use of unsat-
isfactory (occasionally plesiomorphic) characters in phylo-
genetic reconstruction. Various authors have recognized
between five and thirteen families, and strikingly, no two
publications have agreed on the same divisions of the
superfamily into families (Kitching & Rawlins 1998; La-
fontaine & Fibiger 2006). Miller (1991) recognized seven
families: Oenosandridae, Doidae, Notodontidae, Lyman-
triidae, Arctiidae, Aganaidae and Noctuidae. Scoble (1992)
included six families, placing Aganainae as a subfamily
within Noctuidae. Kitching & Rawlins (1998) later recog-
nized three fundamental lineages of Noctuoidea: Oenosan-
dridae, Doidae + Notodontidae, and the quadrifid families
(those where vein MA2 arises very close to, or is stalked
with, MP1 in the forewing, i.e., Arctiidae, Lymantriidae,
Noctuidae, Nolidae and Pantheidae). Most recently, three
landmark publications (Fibiger & Lafontaine 2005; Lafon-
taine & Fibiger 2006; Mitchell
et al.
2006) presented
detailed phylogenies and revised the classification of Noc-
tuoidea three times, each classification having its own limi-
tations and strengths (Roe
et al.
2010). Fibiger &
Lafontaine (2005) proposed a new classification with ten
families: Oenosandridae, Doidae, Notodontidae, Strepsi-
manidae, Nolidae, Lymantriidae, Arctiidae, Erebidae,
Micronoctuidae and Noctuidae. Lafontaine & Fibiger
(2006) proposed a further revision to the classification of
the families of Noctuoidea, in which Nolidae, Strepsiman-
idae, Arctiidae, Lymantriidae and Erebidae
sensu
Fibiger &
Lafontaine (2005) were downgraded to subfamily status
within an expanded family concept of Noctuidae based on
the quadrifid venation of the forewing and the presence of
a tympanal sclerite in the tympanal membrane. In their
view, the superfamily should consist of five families:
Oenosandridae, Doidae, Notodontidae, Micronoctuidae
and Noctuidae.
Within the superfamily, the most controversial family
group taxon is Noctuidae. Many of the traditional subfam-
ilies are now recognized as unnatural (Kitching 1984; Beck
1991, 1992; Lafontaine & Poole 1991; Speidel
et al.
1996;
Kitching & Rawlins 1998; Fibiger & Lafontaine 2005;
Mitchell
et al.
2006). Indeed, the composition and mono-
phyly of many subfamilies is still open to question, and in
particular, the taxonomic composition of the quadrifine
noctuoids (those with a strong vein MA2 in the hindwing)
has remained notoriously difficult to establish. The situa-
tion has been reviewed recently by several authors (Speidel
& Naumann 1995; Fibiger 2003; Ku
¨hne & Speidel 2004;
Holloway 2005, 2008), who have suggested that the mono-
phyly of the group was highly doubtful. At a noctuid
workshop in Denmark in 2002 (Holloway 2005), it was
decided that, prior to any further attempts to redefine
family groups across the superfamily, it was necessary to
gain a clearer understanding of the higher taxonomic
diversity involved by attempting to identify on morpholog-
ical grounds more potentially monophyletic groupings of
genera within the immense diversity of the trifid section of
the superfamily, particularly amongst the much less well
worked quadrifine richness in the tropics. Studies of this
kind would provide a basis for a sampling strategy for
future phylogenetic studies across the group as a whole,
exemplars being selected from significant groupings of
genera and morphologically well-supported concepts of
higher taxa (and subgroups thereof) such as the traditional
Arctiidae and Lymantriidae. This approach was adopted
Molecular phylogeny of Noctuoidea dR. Zahiri et al.
2Zoologica Scripta, 2010 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters

by Holloway (2005) when exploring a broad cross-section
of the Oriental tropical quadrifine fauna from Borneo,
relating it as far as possible to type taxa of available family
group names globally.
Several molecular studies have examined higher level
relationships within the Noctuidae
sensu lato
. Weller
et al.
(1994), using partial sequences of nuclear 28S rRNA
(300 bp) and mitochondrial ND1 (320 bp) from 26 noctu-
oid species, including 10 noctuids, noted that, despite low
levels of support, parsimony analyses consistently grouped
quadrifine noctuids with Arctiidae, and often Lymantrii-
dae, rather than with trifine noctuids (those with vein
MA2 in the hindwing usually vestigial or absent so that
the cubital vein appears to branch into three veins), sug-
gesting paraphyly of Noctuidae. However, vein reduction
in this region also occurs in some Arctiidae and Nolidae
as we discuss later. Subsequent studies based on sequences
of two nuclear genes,
Elongation Factor-1a
(EF-1a) and
Dopa Decarboxylase
(DDC) (Friedlander
et al.
1994; Mitch-
ell
et al.
1997, 2000, 2006; Fang
et al.
2000) provided fur-
ther evidence for the paraphyly of Noctuidae. Mitchell
et al.
(2006) found a strongly supported clade of quadrifine
noctuid moths that included the families Lymantriidae and
Arctiidae. They termed this the L.A.Q. clade (Lymantrii-
dae, Arctiidae and Quadrifine Noctuidae).
Two recent molecular studies on ditrysian Lepidoptera
sampled members of Noctuoidea and found that the enig-
matic family Doidae did not group with the other noctu-
oids, but appeared to be related to Drepanoidea (Regier
et al.
2009; Mutanen
et al.
2010). Otherwise both studies
found Noctuoidea to be monophyletic, with Oenosandri-
dae being sister to the rest and Notodontidae the next
lineage branching off.
However, all these studies had very poor sampling of
the higher taxa putatively belonging to the L.A.Q. clade,
and critically they did not sample type genera of many
higher taxa. Given that the monophyly of many named
groups remains in question, it is crucial to sample the type
genera of each family, subfamily and tribe to assess the
taxonomic limits of a given category.
Previous molecular studies have used only a small num-
ber of molecular markers, usually one to three gene
regions (Wahlberg & Wheat 2008). Here, we present a
phylogenetic hypothesis for higher taxa of Noctuoidea
using new molecular data from eight gene regions.
Materials and methods
We sampled 152 representatives of many major lineages of
the Noctuoidea complex. These comprise four outgroup
taxa and 148 Noctuoidea species representing four families
(Oenosandridae, Notodontidae, Noctuidae and Micro-
noctuidae), 50 subfamilies and 51 tribes, as recognized by
Lafontaine & Fibiger (2006), as well as 16 taxa of uncer-
tain position (Table 1). Based on the results of Regier
et al.
(2009) and Mutanen
et al.
(2010), as well as our own
preliminary analyses, we did not include the family Doi-
dae. We were unable to sample some scarce taxa with
restricted distributions and ⁄or low species richness (e.g.,
subfamilies Cocytiinae, Eucocytiinae and Strepsimaninae
and the type genera of a few tribes ⁄subtribes). To test the
monophyly of the Noctuoidea, we included four species
from three other superfamilies, namely Drepanoidea,
Bombycoidea and Geometroidea. We rooted the clado-
grams with
Thyatira batis
(Drepanidae).
We extracted DNA from one or two legs, dried or
freshly preserved in 96% ethanol, using the DNeasy tissue
extraction kit (QIAGEN, Hilden, Germany). For each
specimen, we sequenced the
cytochrome oxidase subunit I
gene (COI) from the mitochondrial genome, and the EF-
1a, Ribosomal protein S5 (RpS5),
Carbamoylphosphate syn-
thase domain protein
(CAD),
Cytosolic malate dehydrogenase
(MDH),
Glyceraldehyde-3-phosphate dehydrogenase
(GAP-
DH),
Isocitrate dehydrogenase
(IDH) and
wingless
genes,
from the nuclear genome. All genes are protein coding
and have been found to be highly informative for phyloge-
netic analyses at the level of families and superfamilies
(Wahlberg & Wheat 2008; Wahlberg
et al.
2009; Mutanen
et al.
2010). PCR and sequencing protocols follow Wahl-
berg & Wheat (2008). Resulting chromatograms were
checked and DNA sequences aligned by eye using the pro-
gram BioEdit (Hall 1999). Alignment was trivial and the
few insertion ⁄deletion events that were detected, were of
entire codons (in CAD, IDH and RpS5) and could be
easily aligned.
The gene regions were analysed separately and com-
bined in various partitions using parsimony and maximum
likelihood (ML) methods. The data were combined in
three ways: all gene regions together, all nuclear genes
together (i.e., the mitochondrial gene COI excluded) and
all gene regions together with third codon positions
excluded.
Parsimony analyses were undertaken by performing New
Technology heuristic searches in the program TNT (Go-
loboff
et al.
2003). All characters were treated as unordered
and equally weighted. Clade robustness was estimated by
Bremer support (Bremer 1988, 1994) using a script (Pen
˜a
et al.
2006) in TNT. Model-based phylogenetic analyses
were implemented using ML and a GTR+G+I model was
chosen as the most appropriate model of sequence evolution
for each gene partition using FindModel (http://www.
hiv.lanl.gov/content/sequence/findmodel/findmodel.html).
However, we assigned all partitions with the GTR+G
model, as the parameters I (proportion of Invariant posi-
tions) and G (Gamma distribution) are strongly correlated
R. Zahiri et al. dMolecular phylogeny of Noctuoidea
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 2010 3

Table 1 List of taxa with voucher codes and GenBank accession numbers. The families are classified as indicated for Figs 1 and 2
Family Subfamily
Tribe:
subtribe Species Specimen ID
COI-
LCO
COI-
Jerry
EF-1a-
begin
EF-1a-
end Wingless GAPDH RpS5 MDH CAD IDH
Type
status Locality
Outgroup
Drepanidae Thyatirinae
Thyatira batis
MM00027 GU828580 GU828380 GU828919 GU829212 GU829481 GU829743 GU830597 GU830293 GU828083 GU829969 TG ⁄TS Finland
Sphingidae Sphinginae
Sphinx ligustri
NW141-12 EU141358 EU141358 EU136665 EU136665 EU141239 EU141494 EU141391 EU141615 EU141313 EU141550 TG ⁄TS Finland
Bombycidae Bombycinae
Bombyx mori
NW149-1 EU141360 EU141360 EU136667 EU136667 EU141241 EU141495 EU141393 EU141617 EU141315 EU141552 TG ⁄TS USA
Geometridae Archiearinae
Archiearis
parthenias
NW107-1 DQ018928 DQ018928 DQ018899 DQ018899 DQ018869 EU141485 EU141381 EU141604 EU141303 EU141539 TG ⁄TS Sweden
Ingroup
Oenosandridae
Oenosandra
boisduvali
MM07590 GU828791 GU929762 GU829098 GU829377 GU829651 GU829871 GU830751 GU830492 GU828266 GU830173 TG ⁄TS Australia
Oenosandridae
Discophlebia
sp. RZ403 HQ006217 HQ006921 HQ006313 HQ006404 HQ006825 HQ006480 HQ006729 HQ006638 – HQ006551 Australia
Notodontidae Phalerinae
Phalera bucephala
MM00122 GU828607 GU828405 GU828941 GU829235 GU829502 – GU830617 GU830318 GU828108 GU829995 TG ⁄TS Finland
Notodontidae Heterocampinae
Stauropus fagi
MM00981 GU828651 GU828449 GU828983 GU829266 GU829539 GU829780 GU830650 GU830357 GU828148 GU830038 TS Finland
Notodontidae Notodontinae
Notodonta
dromedarius
MM00998 GU828653 GU828451 GU828984 GU829268 GU829540 GU829781§ GU830652 GU830359 GU828150 GU830040 TG ⁄TS Finland
Notodontidae Pygaerinae
Clostera pigra
MM01005 GU828654 GU828452 GU828985 GU829269 GU829541 GU829782 GU830653 GU830360 GU828151 GU830041 Finland
Notodontidae Thaumetopoeinae
Epicoma
melanosticta
MM07592 GU828792 GU929763 GU829099 GU829378 GU829652 GU829872 GU830752 GU830493 GU828267 GU830174 Australia
Notodontidae Thaumetopoeinae
Thaumetopoea
solitaria
MM09888 GU828843 GU929807 GU829144 – GU829692 GU829904 GU830791 GU830534 GU828307 GU830223 TG Greece
Notodontidae Dudusinae
Crinodes besckei
05-srnp-57213 GU828527 – GU828873 GU829175 GU829434 – GU830563 GU830251 GU828039 GU829918 Costa Rica
Notodontidae Nystaleinae
Nystalea striata
05-srnp-4443 GU828525 – GU828871 GU829173 GU829432 GU829717 GU830561 GU830249 GU828037 GU829916 TG Costa Rica
Notodontidae Dioptinae
Scotura
leucophleps
06-srnp-22781 GU828532 GU828334 GU828878 GU829179 GU829439 GU829721 GU830568 GU830256 GU828044 GU829923 Costa Rica
Erebidae Rivulinae
Rivula sericealis
MM01404 GU828664 GU828462 GU828995 GU829278 – GU829791 – GU830370 GU828161 GU830051 TG ⁄TS Finland
Erebidae Boletobiinae
Parascotia
fuliginaria
MM00340 HQ006154 HQ006862 HQ006253 HQ006347 HQ006764 HQ006436 HQ006672 HQ006583 HQ006954 HQ006505 TG ⁄TS Finland
Erebidae Hypenodinae
Hypenodes
humidalis
MM01780 GU828671 GU828469 – GU829285 GU829556 – GU830666 – GU828168 GU830058 TG ⁄TS Finland
Erebidae Hypenodinae
Schrankia
costaestrigalis
RZ27 HQ006192 HQ006896 HQ006288 HQ006382 HQ006800 HQ006461 HQ006705 HQ006613 HQ006987 – Hong Kong
Erebidae Araeopteroninae
Araeopteron
sp. RZ137 HQ006170 HQ006874 HQ006267 HQ006361 HQ006779 – HQ006686 – HQ006966 HQ006515 TG Indonesia
Erebidae Eublemminae Eublemmini
Eublemma
purpurina
RZ7 HQ006237 HQ006940 HQ006332 HQ006424 HQ006845 HQ006491 HQ006748 HQ006655 – HQ006569 TG Hungary
Erebidae Herminiinae
Polypogon
strigilatus
MM01286 GU828663 GU828461 GU828994 GU829277 GU829549 GU829790 GU830660 GU830369 GU828160 GU830050 TG ⁄TS Finland
Erebidae Herminiinae
Paracolax
tristalis
RZ5 HQ006224 HQ006927 HQ006319 HQ006411 HQ006832 – HQ006736 – HQ007013 – Hungary
Erebidae Herminiinae
Herminia
tarsicrinalis
RZ6 HQ006232 HQ006935 HQ006327 HQ006419 HQ006840 HQ006489 ––––TGHungary
Erebidae Herminiinae
Simplicia
pachycera
RZ166 HQ006175 HQ006879 HQ006272 HQ006366 – HQ006448 HQ006691 HQ006599 HQ006971 HQ006520 Ghana
Molecular phylogeny of Noctuoidea dR. Zahiri et al.
4Zoologica Scripta, 2010 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters

Table 1 (Continued)
Family Subfamily
Tribe:
subtribe Species Specimen ID
COI-
LCO
COI-
Jerry
EF-1a-
begin
EF-1a-
end Wingless GAPDH RpS5 MDH CAD IDH
Type
status Locality
Erebidae Scolecocampinae
Scolecocampa
liburna
RZ9 HQ006242 HQ006944 HQ006336 HQ006429 HQ006850 HQ006495 HQ006753 HQ006660 HQ007025 HQ006573 TG USA
Erebidae Hypeninae
Hypena
proboscidalis
MM01545 GU828668 GU828466 GU828999 GU829282 GU829553 GU829794 GU830664 GU830374 GU828165 GU830055 TG ⁄TS Finland
Erebidae Phytometrinae
Phytometra
viridaria
RZ129 HQ006165 HQ006962 HQ006262 HQ006356 HQ006774 HQ006442 HQ006681 HQ006591 HQ006962 HQ006512 TG Finland
Erebidae Phytometrinae
Colobochyla
salicalis
RZ4 HQ006215 HQ006919 HQ006311 HQ006402 HQ006823 HQ006478 HQ006727 HQ006636 HQ007005 – TS Hungary
Erebidae Phytometrinae
Oxycilla ondo
RZ24 HQ006184 HQ006888 HQ006280 HQ006375 HQ006792 HQ006456 – HQ006607 HQ006980 HQ006529 USA
Erebidae Pangraptinae
Pangrapta
bicornuta
RZ40 HQ006216 HQ006920 HQ006312 HQ006403 HQ006824 HQ006479 HQ006728 HQ006637 HQ007006 HQ006550 TG Hong Kong
Erebidae Pangraptinae
P. decoralis
RZ66 HQ006236 HQ006939 HQ006331 HQ006423 HQ006844 – HQ006747 – HQ007022 HQ006568 TG ⁄TS USA
Erebidae Aventiinae
Laspeyria flexula
RZ3 HQ006197 HQ006901 HQ006293 HQ006386 HQ006805 HQ006463 HQ006710 HQ006618 HQ006990 HQ006536 TG ⁄TS Hungary
Erebidae Aventiinae
Corgatha nitens
RZ36 HQ006211 HQ006915 HQ006307 HQ006398 HQ006819 HQ006474 HQ006723 HQ006632 HQ007001 HQ006547 Hong Kong
Erebidae Aventiinae
Prolophota
trigonifera
RZ37 HQ006212 HQ006916 HQ006308 HQ006399 HQ006820 HQ006475 HQ006724 HQ006633 HQ007002 – TS Hong Kong
Erebidae Aventiinae
Metaemene
albiguttata
RZ41 HQ006218 HQ006922 HQ006314 HQ006405 HQ006826 HQ006481 HQ006730 HQ006639 HQ007007 HQ006552 Hong Kong
Erebidae Aventiinae Trisatelini
Trisateles
emortualis
MM04877 GU828707 GU828502 GU829030 GU829319 GU829583 GU829821 GU830695 GU830411 GU828195 GU830093 TG ⁄TS Finland
Erebidae Erebinae Erebini
Erebus ephesperis
RZ11 HQ006161 HQ006866 HQ006258 HQ006353 HQ006770 HQ006440 HQ006677 HQ006587 HQ006959 HQ006510 TG Taiwan
Erebidae Unassigned Anobini
Anoba anguliplaga
RZ332 HQ006206 HQ006910 HQ006302 HQ006395 HQ006814 HQ006469 – HQ006627 – HQ006544 TG Ghana
Erebidae Unassigned Anobini
Marcipa
sp. RZ177 HQ006177 HQ006881 – HQ006368 HQ006785 HQ006450 – HQ006601 HQ006973 HQ006522 Ghana
Erebidae Unassigned
Episparis
group
Masca abactalis
RZ18 HQ006178 HQ006882 HQ006274 HQ006369 HQ006786 HQ006451 HQ006693 – HQ006974 HQ006523 TS Indonesia
Erebidae Unassigned
Ugia insuspecta
RZ45 HQ006221 HQ006925 – HQ006408 HQ006829 HQ006484 HQ006733 HQ006642 HQ007010 HQ006555 Hong Kong
Erebidae Unassigned
Saroba
group
Saroba pustulifera
RZ104 HQ006160 HQ006865 HQ006257 HQ006352 HQ006769 – HQ006676 – – HQ006509 TS Hong Kong
Erebidae Unassigned
Sphingomorpha
chlorea
RZ291 HQ006195 HQ006899 HQ006291 HQ006385 HQ006803 – HQ006708 HQ006616 – – TS Tanzania
Erebidae Unassigned Eulepidotini
Eulepidotis
rectimargo
RZ12 HQ006162 HQ006960 HQ006259 HQ006354 HQ006771 – HQ006678 HQ006588 HQ006960 HQ006511 TG Costa Rica
Erebidae Unassigned Thysaniini
Thysania zenobia
RZ53 HQ006225 HQ006928 HQ006320 HQ006412 HQ006833 HQ006486 HQ006737 HQ006645 HQ007014 HQ006558 TG Costa Rica
Erebidae Unassigned
Oxidercia toxea
RZ295 HQ006196 HQ006900 HQ006292 – HQ006804 – HQ006709 HQ006617 HQ006989 – TS Costa Rica
Erebidae Calpinae Scoliopterygini
Scoliopteryx
libatrix
MM00407 GU828641 GU828439 GU828975 GU829260 GU829532 – GU830643 GU830348 GU828140 GU830028 TG ⁄TS Finland
Erebidae Calpinae Anomini
Anomis involuta
RZ13 HQ006166 HQ006963 HQ006263 HQ006357 HQ006775 – HQ006682 HQ006592 HQ006963 – TG Tanzania
Erebidae Calpinae Anomini
A. metaxantha
RZ55 HQ006227 HQ006930 HQ006322 HQ006414 HQ006835 – HQ006739 HQ006647 HQ007016 HQ006560 TG Taiwan
Erebidae Calpinae Phyllodini
Phyllodes
eyndhovi
RZ56 HQ006228 HQ006931 HQ006323 HQ006415 HQ006836 – HQ006740 HQ006648 – HQ006561 TG Taiwan
Erebidae Calpinae Phyllodini
Miniodes
phaeosoma
RZ153 HQ006173 HQ006877 HQ006270 HQ006364 HQ006782 HQ006446 HQ006689 HQ006597 HQ006969 HQ006518 Ghana
Erebidae Calpinae Calpini
Calyptra thalictri
MM00963 HQ006156 HQ006861 HQ006252 HQ006348 HQ006763 HQ006435 HQ006671 HQ006582 HQ006955 HQ006504 TG ⁄TS Finland
R. Zahiri et al. dMolecular phylogeny of Noctuoidea
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 2010 5

Table 1 (Continued)
Family Subfamily
Tribe:
subtribe Species Specimen ID
COI-
LCO
COI-
Jerry
EF-1a-
begin
EF-1a-
end Wingless GAPDH RpS5 MDH CAD IDH
Type
status Locality
Erebidae Calpinae Calpini
Eudocima
fullonia
RZ16 HQ006174 HQ006878 HQ006271 HQ006365 HQ006783 HQ006447 HQ006690 HQ006598 HQ006970 HQ006519 Malaysia
Erebidae Calpinae Calpini
Plusiodonta
nitissima
RZ333 HQ006207 HQ006911 HQ006303 – HQ006815 HQ006470 HQ006719 HQ006628 – – Costa Rica
Erebidae Calpinae Calpini
Gonodonta uxor
RZ335 HQ006208 HQ006912 HQ006304 – HQ006816 HQ006471 HQ006720 HQ006629 – HQ006545 Costa Rica
Erebidae Calpinae Calpini
Oraesia emarginata
RZ102 HQ006159 HQ006864 HQ006256 HQ006351 HQ006768 HQ006439 HQ006675 HQ006586 HQ006958 HQ006508 TS Hong Kong
Erebidae Calpinae Unassigned
Hypsoropha hormos
RZ17 HQ006176 HQ006880 HQ006273 HQ006367 HQ006784 HQ006449 HQ006692 HQ006600 HQ006972 HQ006521 USA
Erebidae Catocalinae
Serrodes
group
Serrodes campana
RZ318 HQ006202 HQ006906 HQ006298 HQ006391 HQ006810 HQ006467 HQ006715 HQ006623 HQ006995 HQ006540 Taiwan
Erebidae Catocalinae Unassigned
Erygia apicalis
RZ29 HQ006194 HQ006898 HQ006290 HQ006384 HQ006802 – HQ006707 HQ006615 HQ006988 HQ006535 TS Hong Kong
Erebidae Catocalinae Unassigned
Sympis rufibasis
RZ48 HQ006223 – HQ006318 HQ006410 HQ006831 HQ006485 HQ006735 HQ006644 HQ007012 HQ006557 TS Hong Kong
Erebidae Catocalinae Unassigned
Anisoneura salebrosa
RZ38 HQ006213 HQ006917 HQ006309 HQ006400 HQ006821 HQ006476 HQ006725 HQ006634 HQ007003 HQ006548 TS Hong Kong
Erebidae Catocalinae Catocalini
Catocala sponsa
MM04358 GU828700 GU828495 GU829023 GU829312 GU829576 GU829816 GU830688 GU830404 GU828189 GU830086 TG Finland
Erebidae Catocalinae Catocalini
Ulotrichopus macula
RZ241 HQ006185 HQ006889 HQ006281 – HQ006793 HQ006457 HQ006699 HQ006608 – HQ006530 Taiwan
Erebidae Catocalinae Toxocampini
Lygephila pastinum
MM05092 GU828711 GU828506 – GU829323 GU829587 – GU830699 GU830415 GU828199 GU830097 TG Finland
Erebidae Catocalinae Toxocampini
L. maxima
RZ57 HQ006229 HQ006932 HQ006324 HQ006416 HQ006837 HQ006487 HQ006741 HQ006649 – HQ006562 TG Japan
Erebidae Catocalinae Toxocampini
Pantydia diemeni
RZ309 HQ006199 HQ006903 HQ006295 HQ006388 HQ006807 HQ006464 HQ006712 HQ006620 HQ006992 HQ006538 Australia
Erebidae Catocalinae Acantholipini
Acantholipes
circumdata
RZ248 HQ006189 HQ006893 HQ006285 HQ006379 HQ006797 – HQ006702 – HQ006984 HQ006531 TG UAE
Erebidae Catocalinae Acantholipini
A. regularis
RZ135 HQ006168 HQ006872 HQ006265 HQ006359 HQ006777 – HQ006684 – – – TG ⁄TS Russia
Erebidae Catocalinae Melipotini
Melipotis jucunda
RZ58 HQ006230 HQ006933 HQ006325 HQ006417 HQ006838 – HQ006742 HQ006650 HQ007017 HQ006563 TG ⁄TS USA
Erebidae Catocalinae Panopodini
Azeta ceramina
RZ22 HQ006182 HQ006886 HQ006278 HQ006373 HQ006790 – HQ006697 HQ006605 HQ006978 HQ006527 Costa Rica
Erebidae Catocalinae Panopodini
Panopoda rufimargo
RZ59 HQ006231 HQ006934 HQ006326 HQ006418 HQ006839 HQ006488 HQ006743 HQ006651 HQ007018 HQ006564 TG USA
Erebidae Catocalinae Ophiusini
Achaea serva
RZ19 HQ006179 HQ006883 HQ006275 HQ006370 HQ006787 HQ006452 HQ006694 HQ006602 HQ006975 HQ006524 Malaysia
Erebidae Catocalinae Ophiusini
Heteropalpia acrosticta
RZ243 HQ006186 HQ006890 HQ006282 HQ006376 HQ006794 – HQ006700 – HQ006981 – UAE
Erebidae Catocalinae Ophiusini
Ophiusa coronata
RZ21 HQ006181 HQ006885 HQ006277 HQ006372 HQ006789 HQ006454 HQ006696 HQ006604 HQ006977 HQ006526 TG Malaysia
Erebidae Catocalinae Ophiusini
O. tirhaca
RZ246 HQ006187 HQ006891 HQ006283 HQ006377 HQ006795 HQ006458 HQ006701 HQ006609 HQ006982 – TG ⁄TS UAE
Erebidae Catocalinae Ophiusini
Clytie devia
RZ247 HQ006188 HQ006892 HQ006284 HQ006378 HQ006796 HQ006459 – HQ006610 HQ006983 – UAE
Erebidae Catocalinae Pandesmini
Pandesma robusta
RZ321 HQ006204 HQ006908 HQ006300 HQ006393 HQ006812 – HQ006717 HQ006625 HQ006997 HQ006542 TG ⁄TS Spain
Erebidae Catocalinae Ophiusini
Artena dotata
RZ46 HQ006222 HQ006926 HQ006317 HQ006409 HQ006830 – HQ006734 HQ006643 HQ007011 HQ006556 Hong Kong
Erebidae Catocalinae Euclidiini
Mocis latipes
RZ20 HQ006180 HQ006884 HQ006276 HQ006371 HQ006788 HQ006453 HQ006695 HQ006603 HQ006976 HQ006525 Costa Rica
Erebidae Catocalinae Euclidiini
Callistege mi
MM05469 HQ006150 HQ006857 HQ006248 HQ006343 HQ006759 – HQ006667 HQ006578 HQ006950 HQ006500 TS Finland
Erebidae Catocalinae Euclidiini
Euclidia glyphica
RZ82 HQ006239 HQ006942 HQ006333 HQ006426 HQ006847 – HQ006750 HQ006657 HQ007023 HQ006570 TG Finland
Erebidae Catocalinae Audeini
Audea bipunctata
RZ60 HQ006233 HQ006936 HQ006328 HQ006420 HQ006841 – HQ006744 HQ006652 HQ007019 HQ006565 TG ⁄TS Congo
Erebidae Catocalinae Sypnini
Sypnoides fumosa
RZ313 HQ006201 HQ006905 HQ006297 HQ006390 HQ006809 HQ006466 HQ006714 HQ006622 HQ006994 HQ006539 Japan
Erebidae Catocalinae Hypopyrini
Hypopyra capensis
RZ149 HQ006172 HQ006876 HQ006269 HQ006363 HQ006781 – HQ006688 HQ006596 HQ006968 HQ006517 TG Ghana
Erebidae Catocalinae Hulodini
Ericeia subcinerea
RZ39 HQ006214 HQ006918 HQ006310 HQ006401 HQ006822 HQ006477 HQ006726 HQ006635 HQ007004 HQ006549 Hong Kong
Erebidae Catocalinae Hulodini
Hulodes caranea
RZ126 HQ006163 – HQ006260 – HQ006772 – HQ006679 HQ006589 – – TG Malaysia
Erebidae Catocalinae Pericymini
Pericyma cruegeri
RZ99 HQ006244 HQ006946 HQ006338 HQ006431 HQ006852 HQ006497 HQ006755 HQ006662 HQ007027 HQ006575 TG Hong Kong
Erebidae Catocalinae Catephiini
Catephia alchymista
RZ127 HQ006164 HQ006961 HQ006261 HQ006355 HQ006773 HQ006441 HQ006680 HQ006590 HQ006961 – TG ⁄TS Germany
Erebidae Catocalinae Ercheini
Ercheia cyllaria
RZ33 HQ006205 HQ006909 HQ006301 HQ006394 HQ006813 – HQ006718 HQ006626 HQ006998 HQ006543 TG Hong Kong
Erebidae Aganainae
Asota caricae
MM00145 GU828615 GU828413 GU828949 GU829240 GU829509 – GU830624 GU830325 GU828115 GU830003 TG ⁄TS Thailand
Molecular phylogeny of Noctuoidea dR. Zahiri et al.
6Zoologica Scripta, 2010 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters

Table 1 (Continued)
Family Subfamily
Tribe:
subtribe Species Specimen ID
COI-
LCO
COI-
Jerry
EF-1a-
begin
EF-1a-
end Wingless GAPDH RpS5 MDH CAD IDH
Type
status Locality
Erebidae Aganainae
A. heliconia
RZ44 HQ006220 HQ006924 HQ006316 HQ006407 HQ006828 HQ006483 HQ006732 HQ006641 HQ007009 HQ006554 TG Hong Kong
Erebidae Arctiinae Lithosiini
Brunia antica
RZ28 HQ006193 HQ006897 HQ006289 HQ006383 HQ006801 HQ006462 HQ006706 HQ006614 – HQ006534 TS Hong Kong
Erebidae Arctiinae Arctiini:
Ctenuchina
Antichloris viridis
MM05380 HQ006151 HQ006858 HQ006249 HQ006344 HQ006760 HQ006433 HQ006668 HQ006579 HQ006951 HQ006501 Ecuador
Erebidae Arctiinae Arctiini:
Ctenuchina
Ctenucha virginica
AM-94-0396 GU828535 GU828337 GU828881 GU829181 GU829442 GU829722 GU830570 – – GU829926 TG USA
Erebidae Arctiinae Syntomini
Apisa canescens
MM05843 HQ006146 HQ006853 – HQ006339 HQ006765 – HQ006663 – – – TS Oman
Erebidae Arctiinae Syntomini
Syntomis phegea
RZ8 HQ006238 HQ006941 – HQ006425 HQ006846 HQ006492 HQ006749 HQ006656 – – TG ⁄TS Hungary
Erebidae Arctiinae Syntomini
Dysauxes famula
MM00154 GU828619 GU828417 GU828954 GU829244 GU829514 – – GU830328 GU828120 GU830008 Greece
Erebidae Arctiinae Arctiini:
Callimorphina
Coscinia cribraria
MM05671 HQ006149 HQ006856 HQ006247 HQ006342 HQ006758 – HQ006666 – HQ006949 HQ006499 Finland
Erebidae Arctiinae Arctiini:
Callimorphina
Callimorpha
dominula
RZ136 HQ006169 HQ006873 HQ006266 HQ006360 HQ006778 HQ006444 HQ006685 HQ006594 HQ006965 HQ006514 TG ⁄TS Russia
Erebidae Arctiinae Arctiini: Arctiina
Creatonotos
transiens
RZ30 HQ006198 HQ006902 HQ006294 HQ006387 HQ006806 – HQ006711 HQ006619 HQ006991 HQ006537 Hong Kong
Erebidae Arctiinae Arctiini: Arctiina
Arctia caja
MM03713 GU828693 GU828489 – GU829305 GU829573 GU829813 – GU830398 GU828185 GU830080 TG ⁄TS Finland
Erebidae Arctiinae Arctiini: Pericopina
Dysschema
leucophaea
RZ88 HQ006240 – HQ006334 HQ006427 HQ006848 HQ006493 HQ006751 HQ006658 – HQ006571 Costa Rica
Erebidae Arctiinae Unassigned
Pseudophaloe
troetschi
06-srnp-35191 GU828534 GU828336 GU828880 GU829180 GU829441 – GU830569 GU830258 GU828046 GU829925 Costa Rica
Erebidae Lymantriinae Lymantriini
Lymantria
monacha
MM01048 GU828655 GU828453 GU828986 GU829270 GU829542 – GU830654 GU830361 GU828152 GU830042 TG Finland
Erebidae Lymantriinae Leucomini
Leucoma salicis
MM06740 GU828748 GU929722 GU829062 GU829347 GU829611 – GU830719 GU830449 GU828232 GU830132 TG ⁄TS Finland
Erebidae Lymantriinae Nygmiini
Nygmia plana
RZ34 HQ006209 HQ006913 HQ006305 HQ006396 HQ006817 HQ006472 HQ006721 HQ006630 HQ006999 HQ006546 TG Hong Kong
Erebidae Lymantriinae Orgyiini
Orgyia antiqua
RZ130 HQ006167 HQ006964 HQ006264 HQ006358 HQ006776 HQ006443 HQ006683 HQ006593 HQ006964 HQ006513 TG ⁄TS Finland
Erebidae Lymantriinae Arctornithini
Arctornis
sp. RZ89 HQ006241 HQ006943 HQ006335 HQ006428 HQ006849 HQ006494 HQ006752 HQ006659 HQ007024 HQ006572 TG Japan
Erebidae Micronoctuinae Micronoctuini
Micronoctua
sp. RZ138 HQ006171 HQ006875 HQ006268 HQ006362 HQ006780 HQ006445 HQ006687 HQ006595 HQ006967 HQ006516 TG Indonesia
Nolidae Chloephorinae Chloephorini
Psudoips prasinana
MM00107 GU828600 GU828399 GU828934 GU829229 GU829496 GU829754 GU830611 GU830312 GU828101 GU829989 TG Finland
Nolidae Chloephorinae Sarrothripini
Nycteola degenerana
MM00135 GU828612 GU828410 GU828946 GU829238 GU829506 GU829760 GU830621 GU830323 GU828113 GU830000 TG Finland
Nolidae Chloephorinae Sarrothripini
Giaura robusta
RZ31 HQ006200 HQ006904 HQ006296 HQ006389 HQ006808 HQ006465 HQ006713 HQ006621 HQ006993 – Hong Kong
Nolidae Chloephorinae Ariolicini
Paracrama dulcissima
RZ43 HQ006219 HQ006923 HQ006315 HQ006406 HQ006827 HQ006482 HQ006731 HQ006640 HQ007008 HQ006553 TS Hong Kong
Nolidae Chloephorinae Ariolicini
Ariolica argentea
RZ63 HQ006234 HQ006937 HQ006329 HQ006421 HQ006842 – HQ006745 HQ006653 HQ007020 HQ006566 TG Japan
Nolidae Bleninae
Blenina octo
RZ64 HQ006235 HQ006938 HQ006330 HQ006422 HQ006843 HQ006490 HQ006746 HQ006654 HQ007021 HQ006567 TG Sumatra
Nolidae Westermanniinae
Negeta signata
RZ26 HQ006191 HQ006895 HQ006287 HQ006381 HQ006799 HQ006460 HQ006704 HQ006612 HQ006986 HQ006533 Hong Kong
Nolidae Eligminae
Eligma narcissus
RZ97 HQ006243 HQ006945 HQ006337 HQ006430 HQ006851 HQ006496 HQ006754 HQ006661 HQ007026 HQ006574 TG ⁄TS Hong Kong
Nolidae Unassigned
Selepa molybdea
RZ32 HQ006203 HQ006907 HQ006299 HQ006392 HQ006811 HQ006468 HQ006716 HQ006624 HQ006996 HQ006541 Hong Kong
Nolidae Nolinae Nolini
Nola aerugula
MM01776 GU828670 GU828468 GU829001 GU829284 GU829555 – GU830665 GU830376 GU828167 GU830057 TG Finland
Nolidae Eariadinae
Earias clorana
MM06650 GU828747 GU929721 GU829061 GU829346 GU829610 GU829845 GU830718 GU830448 GU828231 GU830131 TG⁄TS Finland
Euteliidae Euteliinae
Eutelia adulatrix
MM00160 GU828621 GU828419 GU828956 GU829246 GU829516 GU829764 GU830629 GU830330 GU828122 GU830010 TG⁄TS Greece
Euteliidae Euteliinae
Marathyssa basalis
RZ23 HQ006183 HQ006887 HQ006279 HQ006374 HQ006791 HQ006455 HQ006698 HQ006606 HQ006979 HQ006528 TS USA
Euteliidae Euteliinae
Targalla subocellata
RZ35 HQ006210 HQ006914 HQ006306 HQ006397 HQ006818 HQ006473 HQ006722 HQ006631 HQ007000 – Hong Kong
R. Zahiri et al. dMolecular phylogeny of Noctuoidea
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 2010 7

Table 1 (Continued)
Family Subfamily
Tribe:
subtribe Species Specimen ID
COI-
LCO
COI-
Jerry
EF-1a-
begin
EF-1a-
end Wingless GAPDH RpS5 MDH CAD IDH
Type
status Locality
Euteliidae Stictopterinae
Lophoptera
hemithyris
MM07614 GU828802 GU929772 GU829107 GU829385 GU829661 GU829879 GU830759 GU830501 GU828274 GU830183 Australia
Noctuidae ?Pantheinae Arcteini
Arcte modesta
RZ54 HQ006226 HQ006929 HQ006321 HQ006413 HQ006834 – HQ006738 HQ006646 HQ007015 HQ006559 TG Malaysia
Noctuidae ?Erebinae Dyopsini
Dyops chromatophila
RZ10 HQ006158 – HQ006255 HQ006350 HQ006767 HQ006438 HQ006674 HQ006585 HQ006957 HQ006507 TG Costa Rica
Noctuidae ?Aediinae
Ecpatia longinqua
RZ25 HQ006190 HQ006894 HQ006286 HQ006380 HQ006798 – HQ006703 HQ006611 HQ006985 HQ006532 Hong Kong
Noctuidae Metoponiinae
Panemeria tenebrata
MM00005 HQ006157 HQ006863 HQ006254 HQ006349 HQ006766 HQ006437 HQ006673 HQ006584 HQ006956 HQ006506 Finland
Noctuidae Acontiinae Acontiini
Acontia lucida
MM00152 GU828617 GU828415 GU828952 GU829243 GU829512 GU829763 GU830627 GU830327 GU828118 GU830006 TG Greece
Noctuidae Acontiinae Acontiini
Emmelia trabealis
MM09893 HQ006147 HQ006854 HQ006245 HQ006340 HQ006756 – HQ006664 HQ006576 HQ006947 – Sardinia
Noctuidae Agaristinae
Periscepta polysticta
MM07669 GU828820 GU929788 GU829125 GU829400 GU829674 GU829892 GU830773 GU830519 GU828289 GU830201 TG ⁄TS Australia
Noctuidae Plusiinae Abrostolini
Abrostola tripartita
MM05132 HQ006152 HQ006859 HQ006250 HQ006345 HQ006761 – HQ006669 HQ006580 HQ006952 HQ006502 TG Finland
Noctuidae Plusiinae Plusiini
Autographa gamma
MM00328 GU828636 GU828434 GU828970 GU829256 GU829528 – GU830640 GU830344 GU828135 GU830023 TG ⁄TS Finland
Noctuidae Amphipyrinae Psaphidini
Brachionycha
nubeculosa
MM01542 GU828667 GU828465 GU828998 GU829281 GU829552 GU829793 GU830663 GU830373 GU828164 GU830054 TS Finland
Noctuidae Amphipyrinae Amphipyrini
Amphipyra
perflua
MM01162 GU828660 GU828458 GU828991 GU829275 GU829546 GU829787 GU830657 GU830366 GU828157 GU830047 TG Finland
Noctuidae Xyleninae Apameini
Apamea crenata
MM01170 GU828661 GU828459 GU828992 GU829276 GU829547 GU829788 GU830658 GU830367 GU828158 GU830048 TG Finland
Noctuidae Xyleninae Caradrinini
Hoplodrina
octogenaria
MM01651 HQ006153 HQ006860 HQ006251 HQ006346 HQ006762 HQ006434 HQ006670 HQ006581 HQ006953 HQ006503 Finland
Noctuidae Xyleninae Xylenini:
Ufeina
Ufeus faunas
RR-98-0914 GU828860 GU929822 GU829163 GU829425 GU829709 GU829911 GU830807 GU830552 GU828320 GU830238 TG USA
Noctuidae Xyleninae Actinotiini
Actinotia polyodon
MM05153 GU828714 GU828509 – GU829326 GU829590 GU829827 GU830702 GU830418 GU828202 GU830100 TG Finland
Noctuidae Bryophilinae
Cryphia raptricula
MM04919 GU828708 GU828503 GU829031 GU829320 GU829584 GU829822 GU830696 GU830412 GU828196 GU830094 Finland
Noctuidae Acronictinae
Acronicta rumicis
MM01529 GU828666 GU828464 GU828997 GU829280 GU829551 GU829792 GU830662 GU830372 GU828163 GU830053 TG Finland
Noctuidae Acronictinae
Craniophora ligustri
MM06745 HQ006148 HQ006855 HQ006246 HQ006341 HQ006757 HQ006432 HQ006665 HQ006577 HQ006948 HQ006498 TS Finland
Noctuidae Raphiinae
Raphia abrupta
CWM-94-0372 GU828548 GU828350 GU828893 GU829193 GU829455 GU829728 GU830579 GU830270 GU828059 GU829939 TG USA
Noctuidae Cuculliinae
Cucullia umbratica
MM04543 GU828701 GU828496 GU829024 GU829313 GU829577 GU829817 GU830689 GU830405 GU828190 GU830087 TG ⁄TS Finland
Noctuidae Pantheinae
Panthea coenobita
MM04583 GU828702 GU828497 GU829025 GU829314 GU829578 – GU830690 GU830406 GU828191 GU830088 TG ⁄TS Finland
Noctuidae Eustrotiinae
Deltote uncula
MM04601 GU828703 GU828498 GU829026 GU829315 GU829579 GU829818 GU830691 GU830407 GU828192 GU830089 Finland
Noctuidae Noctuinae
Noctua fimbriata
MM04752 GU828705 GU828500 GU829028 GU829317 GU829581 GU829820 GU830693 GU830409 GU828194 GU830091 TG Finland
Noctuidae Condicinae
Condica vecors
CWM-95-0471 GU828550 GU828352 GU828895 GU829194 GU829457 – GU830581 – GU828061 GU829941 TG USA
Noctuidae Heliothinae
Pyrrhia umbra
MM05114 GU828712 GU828507 GU829034 GU829324 GU829588 GU829825 GU830700 GU830416 GU828200 GU830098 Finland
Noctuidae Hadeninae Glottulini
Diaphone
capillamentum
MF-05-0053 GU828571 GU828372 GU828913 GU829206 GU829475 GU829738 GU830591 GU830285 GU828076 GU829960 Tanzania
Noctuidae Bagisarinae
Xanthodes albago
MM09894 GU828844 GU929808 GU829145 GU829412 GU829693 – GU830792 GU830535 GU828308 GU830224 Sardinia
–, gene region was not amplified for specimen; TG, type genus; TS, type species; COI, cytochrome oxidase subunit I gene; EF-1a, Elongation Factor-1a; RpS5, Ribosomal protein S5; CAD, Carbamoylphosphate synthase domain
protein; MDH, Cytosolic malate dehydrogenase; GAPDH, Glyceraldehyde-3-phosphate dehydrogenase; IDH, Isocitrate dehydrogenase.
Molecular phylogeny of Noctuoidea dR. Zahiri et al.
8Zoologica Scripta, 2010 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters

and deeply intertwined such that it is impossible to tease
them apart (Ren
et al.
2005; Kelchner & Thomas 2007),
making it likely that complications arise in estimating values
for these parameters. The gamma function is enough for
correcting for the rate variations amongst sites, including
sites which do not change at all in the dataset. ML analyses
were conducted using the web-server RAxML (Stamatakis
et al.
2008). ML bootstrap analysis with 1000 pseudorepli-
cates (Felsenstein 1985) was conducted with RAxML.
Results
Our analyses are based on sequence data from seven
nuclear gene regions (1240 bp of EF-1a, 400 bp of
wing-
less
, 617 bp of RpS5, 850 bp of CAD, 410 bp of MDH,
691 bp of GAPDH and 710 bp of IDH) and one mito-
chondrial gene region (1477 bp of COI), for a total of
6407 aligned nucleotide sites (Table 2). We were not able
to amplify some genes for some taxa (Tables 1 and 2).
The optimal cladograms found by the two methods
(parsimony and ML) for the combined, complete datasets
are very similar, but show novel relationships not previ-
ously suggested (Figs 1 and 2). The monophyly of Noctu-
oidea is strongly supported (BP ‡96; BS ‡18), within
which we find six strongly supported clades that we feel
deserve family status. These are Oenosandridae, Noto-
dontidae, Erebidae, Nolidae, Euteliidae stat. rev. and Noc-
tuidae (Figs 1 and 2). The Notodontidae are found to be
the sister group of all other Noctuoidea, with the Austra-
lian family Oenosandridae branching off next. However,
this pattern of relationships relative to the rest of Noctuoi-
dea is not well supported. Both Oenosandridae and Noto-
dontidae have a trifid forewing venation similar to that of
Geometridae, a character state that appears to be plesio-
morphic relative to the quadrifid forewing venation found
in the other noctuoid families. Relationships amongst the
remaining four families are not clear, although they form
a monophyletic group with very strong support (Fig. 1).
Euteliidae are sister to Noctuidae in ML analyses (Fig. 1),
and sister to the other three families together in parsi-
mony analyses. Similarly, Nolidae are sister to Erebidae in
ML analyses, but form a trichotomy with Erebidae and
Noctuidae in parsimony analyses.
The six strongly supported clades are also found when
only nuclear gene regions are analysed (i.e., the mitochon-
drial gene is excluded) and when third codon positions are
excluded (Supporting information, Appendices S1 and S2).
However, the relationships amongst the clades we are des-
ignating as families (Oenosandridae, Notodontidae, Eut-
eliidae, Erebidae, Nolidae and Noctuidae) are not stable.
The two analyses now place Oenosandridae as sister to the
rest of Noctuoidea, with Notodontidae branching off next,
and this arrangement is quite well supported (BP ‡87 for
the node Notodontidae + the rest) when the third codon
positions are excluded (Supporting information, Appen-
dix S2). The four remaining families always form a well-
supported clade, but their relationships once again vary.
Nuclear gene regions place Nolidae as sister to the rest
and Erebidae as sister to Euteliidae + Noctuidae, whereas
when third codon positions are removed, Euteliidae are
sister to the rest and Erebidae are sister to Noli-
dae + Noctuidae. Of the single gene analyses only CAD
recovers all six family clades as monophyletic (Supporting
information, Appendix S3), although most members of the
clades recovered in the combined analyses do tend to
remain together in a clade with the other genes, and the
non-monophyly of the families is not strongly supported.
As the results of these analyses do not show strongly sup-
ported conflict with the combined, complete analysis, we
henceforth describe in detail only the latter results.
Nolidae come out as a well-supported monophyletic
clade (BP ‡97; BS ‡10) contra Beck (2009). Nolinae (rep-
resented by type genus) are placed as sister to the rest of
the family. Eligminae are confirmed as belonging to this
family, though they lack most of the diagnostic character-
istics listed by Holloway (1998, 2003) and are associated
with
Selepa
, a genus unassigned to any subfamily by Hollo-
way. The subfamily Chloephorinae forms a major clade
that also includes two further groups given subfamily sta-
tus by Holloway: Bleninae, placed as sister to Sarrothri-
pini; and Eariadinae, placed with representatives of
Chloephorini and Ariolicini. The subfamily Westerman-
niinae remains separate; the subfamilies Afridinae, Collo-
meninae and Risobinae were not sampled.
Euteliidae are strongly supported as a monophyletic
group (BP ‡100; BS ‡9). The clade consists of two sub-
families, Stictopterinae and Euteliinae.
Table 2 Basic statistics for the eight gene regions used in this
study
Gene region
Number of
base pairs
Number of
taxa sequenced
Number of parsimony
informative sites
COI 1477 152 607
EF-1a1240 152 386
Wingless 400 146 215
RpS5 617 142 255
GAPDH 691 98 290
CAD 859 127 429
IDH 716 117 337
MDH 407 139 194
Total 6407 1073 2713
COI,
cytochrome oxidase subunit I
gene; EF-1a,
Elongation Factor-1a
; RpS5,
Ribosomal protein
S5
; GAPDH,
Glyceraldehyde-3-phosphate dehydrogenase
; CAD,
Carbamoylphosphate synthase domain
protein; IDH,
Isocitrate dehydrogenase
;
MDH,
Cytosolic malate dehydrogenase
.
R. Zahiri et al. dMolecular phylogeny of Noctuoidea
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 2010 9

Molecular phylogeny of Noctuoidea dR. Zahiri et al.
10 Zoologica Scripta, 2010 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters

The Noctuidae clade, as delimited in Fig. 1, has strong
support (BP ‡99; BS ‡10), the most striking feature of
which is the inclusion of a clade comprising two quadrifine
tribes, Arcteini (represented by type genus) and Dyopsini
(represented by type genus), which were previously
assumed to be related to Erebinae
sensu
Lafontaine &
Fibiger (2006). Otherwise, this clade is made up of trifine
noctuids. The ‘pest clade’ of Mitchell
et al.
(2006) (Helio-
thinae to Xyleninae) has strong BP (100) but weak BS (1)
support.
Our results provide strong support (BP ‡99; BS ‡9) for
the monophyly of Erebidae. Strikingly, many of the basal
divergences in the family show very short branches with
no or low support. Some traditionally recognized families,
subfamilies and tribes show clear evolutionary relation-
ships with strong BS and BP support. For example, there
is clear support for a clade with Pangraptinae (represented
by its type genus) as sister to another well-supported clade
comprising Aganainae (represented by its type
genus) + Herminiinae (represented by its type genus and
three other core herminiines) and Arctiinae (represented
by 12 genera including the type genus) (Fig. 2). The sys-
tematic position of what traditionally has been recognized
as the family Lymantriidae is clearly within Erebidae
(Fig. 2), consistent with the findings of previous molecular
studies.
Oxycilla
, a genus recently excluded from Rivulinae
by Fibiger & Lafontaine (2005), is grouped with
Rivula
(type genus of Rivulinae) with strong support values (BP
‡96; BS ‡20). Both ML and parsimony analyses place the
recently discovered family Micronoctuidae within subfamily
Hypenodinae (BP ‡89; BS ‡14). A strongly supported
clade (BP ‡98; BS ‡18), which we term the boletobiine
clade, places together a large number of taxa with very
diverse feeding habits, including detritivory, fungivory,
lichenivory, frugivory, pod-boring, predation on other
insects and carrion feeding as well as defoliation; associa-
tion of many of these taxa was suggested by Holloway
(2005, 2009).
Our analyses fail to recover some previously recognized
subfamilies within Erebidae as monophyletic groups
(Fig. 2). Both parsimony and ML analyses suggest that
some recent concepts of subfamilies Calpinae, Catocalinae,
Erebinae and Phytometrinae are polyphyletic. For example,
Calpinae
sensu
Lafontaine & Fibiger (2006) consisted of
four tribes, Anomini, Scoliopterygini, Calpini and Phyllo-
dini. Our results placed two of these tribes, Anomini and
Scoliopterygini, into a strongly supported monophyletic
group (BP ‡100; BS ‡23) as in Holloway (2005). This is
well separated from the clade that comprises the five gen-
era of Calpini and two Phyllodini (BP ‡83). Within this
clade, four Calpini (
Calyptra thalictri
, the type genus;
Gon-
odonta
;
Plusiodonta
;
Oraesia
) constitute a monophyletic
group with some support (BP ‡93; BS ‡3), whereas the
fifth calpine,
Eudocima
, was clustered with the two genera
of Phyllodini (
Phyllodes
and
Miniodes
), albeit with low
branch support (BP ‡67; BS ‡1).
Discussion
Until recently, the higher classification of the Noctuoidea
has been based on morphological characters with a pre-
dominantly phenetic approach until the review by Kit-
ching (1984). Higher taxa have been defined on alternative
states of particular adult characters such as ones of wing
venation mentioned earlier, but also: the orientation of the
thoracic tympanum (ventral vs. posterior); the position of
the counter-tympanal hood relative to the spiracle of A1
(anterior vs. posterior); spining of the mid-tibia (present
vs. absent); genitalic structures including everted vesicae
and expanded corpus bursae (Fibiger & Lafontaine 2005).
However, these characters were often found to be in con-
flict, or both states were found to occur in taxa that other-
wise would be considered congeneric, leading to debate
about the relative weight each should be accorded. Genital
characters, including structures of the everted vesica and
within the corpus bursae, have also augmented the body of
morphological information available, providing autapo-
morphies for all or major parts of many higher taxa within
the superfamily, though not significantly so at the family
level, except possibly for Euteliidae and Nolidae. For Ere-
bidae, Fibiger (2003) and Holloway (2005) came to some-
what different conclusions about classification of the
traditional quadrifine noctuids, even though these were
based in part on the same collection of over 2000 genitalia
slides from the Oriental fauna and from the type species
Fig. 1 The phylogenetic hypothesis of the superfamily Noctuoidea based on a maximum likelihood analysis, along with outgroups. Clades
representing families are coloured. Numbers given above branches are bootstrap values (>50%) and numbers below the branch are
Bremer support values for the node. Nodes without Bremer support values do not appear in the most parsimonious trees. The six
families recognized here are indicated. The Erebidae clade (in red) is shown in more detail in Fig. 2. Family group ranking within each
follows from this (i.e., subfamily and below), though assignment of individual taxa reflect previous classifications, particularly Fibiger &
Lafontaine (2005), Lafontaine & Fibiger (2006) and Holloway (2005, 2009). The three taxa transferred in the analysis from the old
‘quadrifine’ noctuid concept to the more restricted one for Noctuidae in the analysis are indicated by asterisks. Names of moths shown in
figure from top to bottom are: Dioptis (Dioptinae), Oenosandra (Oenosandridae), Targalla (Euteliinae), Euchalcia (Plusiinae), Moma
(Acronictinae), Periphanes (Heliothinae), Euxoa (Noctuinae), Eligma (Eligminae) and Hypopyra (Hypopyrini).
R. Zahiri et al. dMolecular phylogeny of Noctuoidea
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 2010 11

of genera on which global family group names were based
(Holloway 2005). There has also been disagreement about
the relative value of larval vs. adult characteristics (e.g.
Beck 2009), although several larval characteristics have
been found to be of significance for higher classification
(Kitching & Rawlins 1998). Besides early stages and adult
characters, there is now another important new set of
characters provided by molecular sequences. The increas-
ing availability of molecular information has brought new
insights into the relationships of noctuoid taxa (Weller
et al.
1994; Mitchell
et al.
1997, 2006), such as the para-
phyly of the old concept of Noctuidae.
Our results, with more molecular data than have been
used previously for this group of moths, point to six well-
supported major lineages that we have defined as families.
Two of the major lineages are well-recognized taxa that
have often been considered families within Noctuoidea,
i.e., Oenosandridae and Notodontidae. Oenosandridae are
a small family, only known from Australia, comprising
eight species in four genera (Nielsen
et al.
1996), which
mainly feed on Myrtaceae (Miller 1991). Miller (1991)
excluded
Oenosandra
from Notodontidae based on the
non-homology of scale tuft structures on the female ter-
gite 7 (the hairs are used to cover the egg masses), placing
it in a separate family. Notodontidae contain approxi-
mately 4200 species and occur worldwide. The other four
lineages have been split into as many as 10 families, with
arctiines, lymantriines and nolines frequently being consid-
ered to be sufficiently distinct from the rest to warrant full
family status.
Wing venation has been thought to be informative of
the phylogenetic relationships of noctuoids, with the quad-
rifine hindwing defining a group comprising our Nolidae,
Erebidae and Euteliidae and the trifine hindwing defining
Noctuidae. However, our results suggest that trifine moths
have evolved from quadrifines multiple times, e.g.,
amongst some Erebidae, such as Arctiinae: Syntomini
(Griveaud 1964; Holloway 1988), and Nolidae (Holloway
1998, 2003) show hindwing vein reduction in addition to
Fig. 2 Continuation of phylogram in Fig. 1. The phylogenetic
hypothesis of the family Erebidae based on a maximum likelihood
analysis. Numbers given above branches are bootstrap values
(>50%) and numbers below the branch are Bremer support values
for the node. Nodes without Bremer support values do not appear
in the most parsimonious trees. Names of moths shown in figure
from top to bottom are: Scoliopteryx (Scoliopterygini), Lymantria
(Lymantriinae), Peridrome (Aganainae), Syntomis (Syntomini),
Euplagia (Arctiinae), Eudocima (Calpinae), Calyptra (Calpinae),
Oruza (Aventiinae), Eublemma (Eublemminae), Sypna (Sypnini),
Erebus (Erebinae), Spirama (Hypopyrini) and Catocala
(Catocalinae).
Molecular phylogeny of Noctuoidea dR. Zahiri et al.
12 Zoologica Scripta, 2010 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters

the Noctuidae sensu stricto. Furthermore, the tribes Arc-
teini and Dyopsini, previously classified as quadrifines, are
part of a well-supported clade otherwise consisting of tri-
fines (some of the more basal ‘trifine’ subfamilies, e.g.,
Plusiinae, Pantheinae, Bagisarinae also have a quadrifine
hindwing). The placement of
Arcte
is corroborated by
morphological characters (see Holloway 2005, 2009),
whereas
Dyops
has not yet been investigated morphologi-
cally in this context.
We follow Fibiger & Lafontaine (2005) in employing
the family name Erebidae, which was previously used as a
subfamily name by Forbes (1954). The family group
names Arctiidae, Herminiidae and Erebidae were
described in the same publication (Leach 1815) and thus
have equal priority, and all three have priority over Cato-
calidae. However, both Arctiinae and Herminiinae are
well-known taxa generally considered to be subfamily level
(Holloway 2008) or family level (e.g. Kitching 1984) taxa,
and thus we consider that it would generate less confusion
to adopt the name Erebidae for the larger group of noc-
tuoids that includes both Arctiinae and Herminiinae.
Within Erebidae, relationships of only a few lineages
are well supported. The monophyly of Aganainae + He-
rminiinae + Arctiinae clade is strongly supported by our
analyses. The association of Herminiinae (Renia) with arc-
tiines was found in an earlier morphological study by
Jacobson & Weller (2002). The clade also has a morpho-
logical synapomorphy in the prespiracular position of the
counter-tympanal hood. Adults of many aganaines and
arctiines are visually striking and aposematic, and agana-
ines and herminiines share long labial palps and a bare
lower frons. Kitching (1984) was the first to exclude
Aganainae and Herminiinae from Noctuidae based on the
prespiracular hood, which was then thought to be plesio-
morphic. In addition, the clade may be characterized by
two further synapomorphies: modified foretibia in the
males of most genera and a swollen metepimeron ventral
to pocket IV (Kitching 1984). Holloway (2008) also dis-
cussed this grouping, commenting on biological differ-
ences between the groups, Herminiinae being generally
cryptic, feeding on vegetable detritus, and Aganainae being
aposematic, feeding on the same suite of toxic cardeno-
lide-synthesizing plant families (Apocynaceae, Asclepiada-
ceae and Moraceae) as the danaine Nymphalidae and
other moth genera such as
Glyphodes
(Crambidae) and
Agathia
(Geometridae). We have also found that Pangrap-
tinae +
Masca
, a member of the
Episparis
group of genera
of Holloway (2005), are sister to the Aganainae + Hermi-
niinae + Arctiinae clade with good support. Pangraptinae
were previously thought to be associated with Eublem-
minae (Fibiger & Lafontaine 2005), but see Holloway
(2005, 2008, 2009) and below.
The boletobiine clade (Fig. 2) has several potential mor-
phological synapomorphies. Many of the included taxa
have the central part of the valve heavily sclerotized with
several processes arranged transversely, with the cucullus
reduced to a membranous flap. In addition, the female
genitalia often have a ring of claw-like spines in the corpus
bursae, the base of which is frequently narrow and coiled
to some degree. The larvae have the first two pairs of pro-
legs reduced and are often warty or pubescent. They feed
on a diversity of resources as mentioned above. Much of
this clade is made up of a revised and enlarged concept of
Aventiinae, together with Eublemminae (Holloway 2009),
as well as many Araeopteroninae and Phytometrinae. We
identify the clade as boletobiine from the oldest family
group name amongst the included taxa (Boletobiinae).
Phylogenetic relationships within the clade are in much
need of study, and promise to reveal considerable insight
into morphological and life history evolution in quadri-
fines.
The separation in the molecular results of the calpine
clade from the scoliopterygine clade and the inclusion of
Phyllodini in the former provides an object lesson on how
the sharing of peculiar morphological adaptations may
mislead in classification, and how shared features of a
more subtle nature may be overlooked in an unchallenged
traditional classification. A robust, highly developed fruit-
piercing (and in some cases skin piercing and blood suck-
ing) proboscis is found widely in Calpini and in some
more robust scoliopterygines such as
Anomis
, with many
similar features in the structure. But the molecular results
point to this being a homoplasy.
Phyllodini, with a spined adult mid-tibia, were tradition-
ally catocalines, and Calpini, with an unspined mid-tibia,
were traditional ophiderines, indeed including the type
genus thereof. The grouping of Phyllodini with
Eudocima
in the molecular analysis brings into greater prominence
several features shared by these groups (Holloway 2005):
leaf mimicry in the forewing and flash colouration in the
hindwing of the adults; Menispermaceae as a favoured lar-
val host family; a similar method of pupation in a leaf
shelter.
The Euteliidae lineage is undoubtedly a well-defined
group of noctuoids and is raised here to family level, thus
supporting the findings of Mitchell
et al.
(2006). Kitching
(1987) demonstrated that Euteliinae and Stictopterinae are
related and form a monophyletic group, based on a large
number of synapomorphies including: reduced female
frenulum, modified basiconic sensilla on the proboscis,
presence of a small oval plate in the ductus ejaculatorius,
anal papillae modified so that their inner surfaces are
directed posteriorly and the counter-tympanal hood has a
unique double structure (Richards 1932; Holloway 1985;
R. Zahiri et al. dMolecular phylogeny of Noctuoidea
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 2010 13

Kitching 1987; Kitching & Rawlins 1998). The host plant
range of the two subfamilies embraces mostly lactiferous
plants, with Euteliinae favouring Anacardiaceae and Stic-
topterinae having distinct lineages on Calophylla-
ceae ⁄Clusiaceae, Dipterocarpaceae and Euphorbiaceae
(Holloway 1985).
The relationships of the six clades remain somewhat
ambiguous, although it is clear that Euteliidae, Nolidae,
Erebidae and Noctuidae together form a monophyletic
group. The position of Oenosandridae varies in our ana-
lyses, with the nuclear only and third codon position
excluded analyses placing it as sister to the rest of Noctu-
oidea. This position has also been found in the two recent
studies on Ditrysia (Regier
et al.
2009; Mutanen
et al.
2010). Our full, combined dataset places Notodontidae in
this position, with Oenosandridae diverging next, but this
is not well supported. It appears that all poorly supported,
unstable relationships are characterized by short basal
branches, especially within Erebidae (Fig. 2). Such patterns
of short branches and low support values have been
thought to indicate rapid radiations (Tajima 1983; Wiens
et al.
2008; Kodandaramaiah
et al.
2010).
Elucidating the evolutionary history of the massive Ere-
bidae clade (potentially including 40 000 species) will
require more intensive sampling. Further studies are also
needed to identify the reasons for the short basal branches,
i.e., whether there is a historical explanation behind them
(e.g., rapid radiation), or whether it is simply an artefact of
insufficient data. Noctuoidea represent a unique oppor-
tunity to investigate the reasons underlying massive diver-
sification in phytophagous insects, and a first step in such
investigations is the identification of monophyletic groups
and their interrelationships.
The sampling strategy that we have adopted, described
in the introduction, is being used to establish priorities for
ongoing studies, to test further the robustness of the four
major clades, both in relation to each other and internally.
Work in progress on an increased sample of taxa, parti-
cularly in Erebidae, shows that the clades remain distinct
and robust, and that ‘known unknowns’, when included,
fall within these clades and tend to reinforce the well sup-
ported parts of their internal structure, rather than per-
turbing it (R. Zahiri
et al.
unpublished data).
In summary, we have shown that there are six strongly
supported lineages in Noctuoidea that can be assigned
family status. Four additional groups that we were unable
to sample, Dilobinae, Cocytiinae, Eucocytiinae and
Strepsimaninae, may prove to be further independent lin-
eages. There are also a few genera of a similar nature that
appear to be noctuoid but have yet to be assigned to a
family, such as
Kenguichardia, Plagerepne
and
Clytophylla
(Holloway
et al.
2001; Holloway 2003). Within the newly
circumscribed families, Erebidae require much more atten-
tion; many traditional subfamilies and tribes (e.g. Calpinae
and Chloephorinae of previous authors) were found to be
polyphyletic, even with our limited sampling. The conclu-
sion that the Euteliidae are not closely related to Erebidae
was surprising and also needs further investigation.
Acknowledgments
This work has been financially supported by the Academy
of Finland (grant nos. 118369 and 129811) awarded to
NW, (grant no. 1110906) awarded to LK, and
CIMO + Finnish Cultural Foundation awarded to RZ.
The main source of samples are the LepTree project
headed by Charles Mitter
et al.
(US NSF award
#0531769); Daniel H. Janzen (US NSF #DEB0072730
and DEB0515699); Royal Museum for Central Africa, Bel-
gium (Ugo Dall’Asta); Roger C. Kendrick (Kadoorie Farm
and Botanic Garden, Hong Kong); Zoologisch Museum
Amsterdam (Rob de Vos); Natural History Naturalis, Lei-
den (Erik J. van Nieukerken); Szabolcs Sa
´fia
´n (Hungary);
Hungarian Natural History Museum (Laszlo Ronkay);
Zoological Institute of the Russian Academy of Sciences,
St. Petersburg (Alexej Matov); Michael Fibiger (Sorø,
Denmark); Natural History Museum, University of Oslo,
Norway (Leif Aarvik); Jo
¨rg-Uwe Meineke (Germany) and
Shen-Horn Yen (Taiwan). The authors also acknowledge
the Natural History Museum London, the Malaysian Nat-
ure Society, the Swedish Museum of Natural History and
Len Willan (CSIRO Entomology) for the permission to
use the moth images depicted in the tree. We thank
Charles Mitter, David Wagner and an anonymous referee
for detailed comments on previous versions of the manu-
script.
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Supporting Information
Additional Supporting Information may be found in the
online version of this article:
Appendix S1. Phylogenetic tree from the maximum
likelihood analysis of the all nuclear genes data set (the
mitochondrial gene COI excluded). Support values are
bootstrap values.
Appendix S2. Phylogenetic tree from the maximum like-
lihood analysis of the all gene regions together third codon
positions excluded. Support values are bootstrap values.
Appendix S3. Phylogenetic trees from the maximum
likelihood analysis of the single gene analyses. Support val-
ues are bootstrap values.
Please note: Wiley-Blackwell are not responsible for the
content or functionality of any supporting materials sup-
plied by the authors. Any queries (other than missing
material) should be directed to the corresponding author
for the article.
Molecular phylogeny of Noctuoidea dR. Zahiri et al.
16 Zoologica Scripta, 2010 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters