TURUN YLIOPISTON JULKAISUJA
ANNALES UNIVERSITATIS TURKUENSISSARJA - SER. AII OSA - TOM. 268
BIOLOGICA - GEOGRAPHICA - GEOLOGICA

Molecular SySteMaticSof Noctuoidea (iNSecta, lepidoptera)reza zahiri

TURUN YLIOPISTO
UNIVERSITY OF TURKU
Turku 2012

From the Laboratory of Genetics, Division of Genetics and Physiology, Department of
Biology, University of Turku, FIN-20012, Finland
Supervised by:Docent Niklas Wahlberg
University of Turku
FinlandCo-advised by:
Ph.D. J. Donald Lafontaine
Canadian National Collection of Insects, Arachnids and Nematodes
CanadaPh.D. Ian J. Kitching
Natural History Museum
U.K.
Ph.D. Jeremy D. Holloway
Natural History Museum
U.K.
Reviewed by:Professor Charles Mitter
University of Maryland
U.S.A.
Dr. Tommi Nyman
University of Eastern Finland
Finland
Examined by:
Dr. Erik J. van Nieukerken
Netherlands Centre for Biodiversity Naturalis, Leiden
The Netherlands
Cover image: phylogenetic tree of Noctuoidea
ISBN 978-951-29-5014-0 (PRINT)
ISBN 978-951-29-5015-7 (PDF)
ISSN 0082-6979
Painosalama Oy – Turku, Finland 2012

To Maryam, my mother and father

MOLECULAR SYSTEMATICS OF NOCTUOIDEA (INSECTA, LEPIDOPTERA)
Reza ZahiriThis thesis is based on the following original research contributions, which are referred toin the text by their Roman numerals:

I

Zahiri, R, Kitching, IJ, Lafontaine, JD, Mutanen, M, Kaila, L, Holloway,
JD & Wahlberg, N (2011) A new molecular phylogeny offers hope for astable family-level classification of the Noctuoidea (Lepidoptera).
Zoologica Scripta, 40, 158–173

II

Zahiri, R, Holloway, JD, Kitching, IJ, Lafontaine, JD, Mutanen, M &Wahlberg, N (2012) Molecular phylogenetics of Erebidae (Lepidoptera,
Noctuoidea). Systematic Entomology, 37,102–124

III

Zaspel, JM, Zahiri, R, Hoy, MA, Janzen, D, Weller, SJ & Wahlberg, N(2012) A molecular phylogenetic analysis of the vampire moths and theirfruit-piercing relatives (Lepidoptera: Erebidae: Calpinae). Submitted
manuscriptIV

Zahiri, R, Lafontaine, JD, Holloway, JD, Kitching, IJ, Schmidt, BC, Kaila,
L, & Wahlberg, N. Major lineages of Nolidae (Lepidoptera, Noctuoidea)
elucidated by molecular phylogenetics. Submitted manuscript

V

Zahiri, R, Lafontaine, JD, Kitching, IJ, Holloway, JD, Schmidt, BC,
Mutanen, M & Wahlberg, N. Relationships of the early lineages ofNoctuidae (Lepidoptera, Noctuoidea) based on eight gene regions.
ManuscriptAbstract
In this thesis, I conduct a series of molecular systematic studies on the large phytophagous
moth superfamily Noctuoidea (Insecta, Lepidoptera) to clarify deep divergences and evolutionary
affinities of the group, based on material from every zoogeographic region of the globe.
Noctuoidea are the most speciose radiations of butterflies and moths on earth, comprising about aquarter of all lepidopteran diversity. The general aim of these studies was to apply suitably
conservative genetic markers (DNA sequences of mitochondrial—mtDNA—and nuclear gene—
nDNA—regions) to reconstruct, as the initial step, a robust skeleton phylogenetic hypothesis forthe superfamily, then build up robust phylogenetic frameworks for those circumscribedmonophyletic entities (i.e., families), as well as clarifying the internal classification ofmonophyletic lineages (subfamilies and tribes), to develop an understanding of the major lineages
at various taxonomic levels within the superfamily Noctuoidea, and their inter-relationships. The
approaches applied included: i) stabilizing a robust family-level classification for the superfamily;ii) resolving the phylogeny of the most speciose radiation of Noctuoidea: the family Erebidae; iii)
reconstruction of ancestral feeding behaviors and evolution of the vampire moths (Erebidae,
Calpinae); iv) elucidating the evolutionary relationships within the family Nolidae and v)
clarifying the basal lineages of Noctuidae sensu stricto. Thus, in this thesis I present a wellresolved molecular phylogenetic hypothesis for higher taxa of Noctuoidea consisting of six
strongly supported families: Oenosandridae, Notodontidae, Euteliidae, Erebidae, Nolidae, andNoctuidae. The studies in my thesis highlight the importance of molecular data in systematic andphylogenetic studies, in particular DNA sequences of nuclear genes, and an extensive samplingstrategy to include representatives of all known major lineages of entire world fauna of Noctuoidea
from every biogeographic region. This is crucial, especially when the model organism is as
species-rich, highly diverse, cosmopolitan and heterogeneous as the Noctuoidea, traits that
represent obstacles to the use of morphology at this taxonomic level.

CONTENTS
CONTENTS .......................................................................................................................6
1. INTRODUCTION .........................................................................................................7
1.1
1.2
1.3
1.4

Why Noctuoidea? ....................................................................................................7
Status of Noctuoidea................................................................................................8
Initiation of molecular phylogenetic approaches ................................................... 10
Outline of the thesis ............................................................................................... 11

2. MATERIAL AND METHODS .................................................................................. 13
2.1 Sampling strategy .................................................................................................. 13
2.2 Molecular markers ................................................................................................. 13
2.3 Phylogenetic analyses and character optimizations ............................................... 14
3. RESULTS AND DISCUSSION .................................................................................. 16
3.1
3.2
3.3
3.4
3.5

Phylogenetic hypothesis for Noctuoidea ............................................................... 16
Pattern of relationships among major lineages of Noctuoidea .............................. 17
Phylogenetic hypothesis for quadrifid Noctuoidea................................................ 19
Character optimizations ......................................................................................... 21
Evolution of host-plant associations in Noctuoidea .............................................. 22

4. CONCLUSIONS AND FUTURE DIRECTIONS..................................................... 29
4.1 Conclusions .............................................................................................................. 29
4.2 Future directions ....................................................................................................... 30
5. ACKNOWLEDGEMENTS ........................................................................................ 32
6. REFERENCES ............................................................................................................ 34
APPENDIX ......................................................................................................................37
ORIGINAL PUBLICATIONS ....................................................................................... 49

Introduction

7“In scientific investigations, it is permitted to invent any hypothesis and, if it explainsvarious large and independent classes of facts, it rises to the rank of a well-groundedtheory”
Charles Darwin

1. INTRODUCTION
1.1 Why Noctuoidea?
I quote a magnificent passage fromCharles Darwin who said ‘‘all the organic
beings which have ever lived on this earth
have descended from some one primordialform.’’ It can be obviously interpreted
from this simple and meaningful passage
that every characteristic of every species
on Earth is the outcome of an evolutionary
history (Futuyma, 2005). The evolutionary
perspective and phylogenetic relationships
have illuminated every subject in biology,
from the molecular and morphology levelto ecosystem and beyond. The geneticist
Theodosius Dobzhansky (1973) famously
argued that ‘‘Nothing in biology makes
sense, except in the light of evolution.’’Evolution governs diversity on Earth, andinsects are the most diverse organisms in
the whole history of life (Grimaldi &Engel, 2005). Consequently, insects should
provide profound insights into evolution.
The Order Lepidoptera (moths andbutterflies) is one of four super-radiationsof insects (along with beetles, flies andwasps) that account for the majority ofanimal life on Earth. Noctuoidea are thelargest superfamily within Lepidoptera—
belonging to a large ditrysian clade that
also includes e.g., geometroids andbombycoids (Regier et al., 2009, Mutanen
et al., 2010)—with approximately 45,000
described (Nieukerken et al., 2011) andmany unknown as well as unnamed
species, particularly from tropical regions.
To understand Noctuoidea evolution,
theirsystemsofevolutionary
relationships—phylogenies
basedon
extensive evidence from living lineages ofnoctuoids—mustberecognized.
Fortunately, the monophyly of Noctuoidea
is firmly established. It is based on thepresence of a single apomorphic character,the metathoracic tympanal organ (Miller,
1991). This organ is a highly specialized
hearing apparatus that detects theecholocation signals of bats (Kitching &Rawlins, 1998); however, there isincreasing evidence that the tympanum
may also be involved in reception ofmating signals (Kitching & Rawlins,1998).
Noctuoidea, like most lepidopterans,
are plant feeding as caterpillars and nectar
feeding as adults, and they are a prominentelementofterrestrial
ecosystems,
functioning as herbivores, pollinators andprey, as well as being one of the mostdamaging groups of pests to agriculture(Regier et al., 2009). Of the approximately
6,000 Lepidoptera species noted to be of
economic importance by Zhang (1994),
about one-quarter belong to Noctuoidea.
Although a large number of these can be
assigned to what Mitchell et al. (2006)
termed the ‘pest clade’, many more are8Introduction

distributed across the whole superfamily
Noctuoidea in over 500 genera (Zhang,1994). The caterpillars of many noctuoid
genera have massive economic impact
annually (Kitching, 1984). In addition, theadults of some genera damage fruit cropsby piercing the skins to suck juices(Baenziger, 1982).
The study of relationships among major
lineagesoforganisms—phylogenetic
analysis—has been closely associated withthe classification and naming of thediversity of life on Earth (i.e., taxonomy),which both are a branch of the science ofsystematics.
The
classificationoforganisms has been one of the major
ongoing accomplishments of humansociety (Wilson, 2000), but is still far fromcompletion, both in terms of the inventoryof species and of the classification of thosespecies in a hierarchical system that has aphylogenetic basis. However, there has
been a striking improvement in the theory
and computational methodology for
inferring phylogenies (Regier et al., 1995).
In particular, the use of molecular data, inparticular DNA sequences, is becoming
increasingly important for testing andimproving classifications, especially forhighly diverse groups of organisms such as
insects.
The main theme of my thesis, as well as
my ongoing research, is to improve ourunderstanding of the reasons and causes
behind the flourishing diversification of
Noctuoidea on Earth. To address tens ofsuchquestions—diversityof
theiradaptations, biomass, species-richness,
ecological and economical impacts, etc.—
it is necessary, as a first step, to place
Noctuoidea and its major lineages in aphylogenetic context, by reconstructing astrong phylogenetic hypothesis. The
resolution of a stable, extrapolative higherlevel classificatory structure for the major
lineages of Noctuoidea, and understanding
their phylogenetic relationships, is also ofimportance for pest bionomic studies.
1.2 Status of Noctuoidea
Historically, the classification of
noctuoid moths has been highly unstable,with different classification systems being
used by different authors. It seems that thefundamental distinction between the
different systems is based on the use of
unsatisfactory(occasionallyplesiomorphic) characters in phylogenetic
reconstruction. Various authors haverecognized between five and thirteen
families, and strikingly, no two
publications have agreed on the samedivisions of the superfamily into families(Kitching & Rawlins, 1998, Lafontaine &Fibiger, 2006). For instance, in Figure 1, I
have summarized most recent Noctuoidea
classifications and have compared themwith the most recent one, which ispresented in this thesis (I). Kitching (1984)
published a historical review of noctuid
subfamily relationships and showed thatthe higher classifications of Noctuidae
used up to that time had been based uponsuperficial resemblance and vaguely
defined characters, rather than on rigorous
applicationofcladisticprinciples.
Subsequently, Speidel and co-workers
attempted to progress beyond the age of
traditional
morphologicalnoctuidtaxonomy by initiating investigations
based mainly on the male genitalia and the
tympanal region, which they consideredparticularly useful in elucidating the basic
relationships of the noctuid subfamilies(Speidel & Naumann, 1995, Speidel et al.,
1996, Kühne & Speidel, 2004, 2005).

Introduction

9

Figure 1 Different classification systems of the superfamily Noctuoidea that have been used since Kitching(1984) to date (I). In every classification moths (except the one for Micronoctuidae for which the author’s
portrait—the late Michael Fibiger—is used) indicate family-group name being used in the system.

Their work set the stage for the study ofthe systematics of noctuoids, and made it
clear that increased character and taxon
sampling were necessary to resolve therelationships of the diverse clades.
Recently, three landmark publications(Fibiger & Lafontaine, 2005, Lafontaine &Fibiger, 2006, Mitchell et al., 2006)
presented detailed phylogenetic hypotheses
and revised the classification of
Noctuoidea three times (Figure 1), eachclassification having its own limitations
and strengths (Roe et al., 2010). Fibiger &Lafontaine (2005) proposed a rather newclassification
withtenfamilies:
Oenosandridae, Doidae, Notodontidae,
Strepsimanidae, Nolidae, Lymantriidae,
Arctiidae, Erebidae, Micronoctuidae andNoctuidae (Figure 1). Later on, Lafontaine& Fibiger (2006) proposed a further
revision to the classification of the families
of Noctuoidea, in which Nolidae,
Strepsimanidae, Arctiidae, Lymantriidae

10Introduction

and Erebidae sensu Fibiger & Lafontaine(2005) were downgraded to subfamily
status within an expanded family conceptof Noctuidae based on the quadrifid
venation of the forewing and the presenceof a tympanal sclerite in the tympanalmembrane. In their view, the superfamily
shouldconsistoffivefamilies:
Oenosandridae, Doidae, Notodontidae,
Micronoctuidae and Noctuidae (Figure 1).
To resolve the dominant complexity ofrelationships in higher systematics ofNoctuoidea, it was crucial to understandtheir phylogenetic relationships bybuilding a robust skeleton phylogenetic
hypothesis, i.e., a phylogeny that was
based on certain specialized features, andthat had a common ancestor and uniqueevolutionary history. To achieve a morerobust phylogenetic hypothesis, one
strategy is to increase the number ofcharacters to obtain a dataset with a strong
phylogenetic
signal.
In
molecularsystematics, datasets with a weak
phylogenetic ‘signal’ tend to be strongly
influenced by the assumptions made by the
analytical methods applied, whereas
datasets with a strong phylogenetic signal
are not influenced as much (Wahlberg &Wheat, 2008). One avenue for acquiring
more characters is to use morphology.
However, a species-rich, cosmopolitan andheterogeneous group such as theNoctuoidea (Speidel & Naumann, 1995)
with a vast number of species presents
obstacles to the use of morphology at thistaxonomic level. The adults and larvae of
species in Noctuoidea exhibit abewildering diversity of size, coloration,
adaptation, behaviour and ecology(Kitching&
Rawlins,
1998).
Morphological data are thus often difficult
to homologize and code, require great
experience to identify character statescorrectly and can be subject to extensive
homoplasy (character convergence andreversal). As a result, morphologicalanalyses have often failed to determine
relationships among most groups withconfidence.
1.3 Initiation of molecular phylogenetic
approaches
As noted above, until recently, thehigher systematics of Noctuoidea had been
based
primarilyonmorphologicalcharacters with a predominantly phenetic
approach, until the introduction andapplication of cladistic philosophy by
Kitching (1984, 1987) and Miller (1991).
More recently, molecular data, inparticularDNA
sequencesofmitochondrial and nuclear genes (mtDNA
and nDNA, respectively), have opened
new and fruitful avenues for the study ofphylogenetic relationships. Molecularstudies have often been based on a small
number of molecular markers, usually
between one and three genes (Sperling,
2003, Wahlberg & Wheat, 2008). The
utility of using many mitochondrial genes
alone is certainly questionable, as they
have a shared evolutionary history, andeven entire mitochondrial genomes(15,000–20,000 bp in insects) fail toprovide robust inferences at deep levels(Cameron et al., 2004). However, they areuseful in the recognition of cryptic species
and sometimes resolve relationships
among closely related species and genera(Lafontaine & Schmidt, 2010). They have
proven most valuable for relatively recent
divergences, especially those of midTertiary and later age, although they havealso been applied, albeit infrequently, atdeeper levels (Wiegmann et al., 2000). In
contrast, protein-coding nuclear genes
generally have slower mutation rates
compared with mtDNA. They are mostIntroduction
frequently used to study older evolutionary
divergences and are particularly good atresolving deeper nodes in phylogenetic
hypotheses, where they have been
important in establishing the family,
subfamily and tribal classification of
Lepidoptera (Regier et al., 2009, Wahlberg
et al., 2009, Mutanen et al., 2010). Overthe past few years, a series of papers havebeen published with a shared objective: tocontribute to the development of a moresatisfactory classification of Noctuoidea atlevels above that of the genus andparticularly the ‘quadrifid’ part—those
noctuoids with forewing vein M2 arises
closer to the origin of M3 than M1, in thelower part of the discal cell—of thesuperfamily. It began with the explorationof molecular markers by Weller et al.
(1994), who were then followed by
Mitchell et al. (1997, 2000, 2006). The
utility of DNA sequences was undeniable,
and systematists were able to gain
fascinating insights that were not obvious
before, e.g., the polyphyly of the oldconcept of Noctuidae (Mitchell et al.,1997). Mitchell et al. (2006) found astrongly supported clade of quadrifine
noctuid moths that also included thefamilies Lymantriidae and Arctiidae. They
termedthis
the
L.A.Q.
clade(Lymantriidae, Arctiidae and Quadrifine
Noctuidae).
Two recent molecular studies on
ditrysian Lepidoptera sampled members of
Noctuoidea and found that the enigmaticfamily Doidae did not group with the othernoctuoids, but instead appeared to berelated to Drepanoidea (Regier et al., 2009,
Mutanen et al., 2010). Otherwise bothstudies found Noctuoidea to bemonophyletic, with Oenosandridae being
sister to the rest and Notodontidae the next
lineage branching off.

11

However, all of these studies had very
poor sampling of the higher taxa putatively
belonging to the L.A.Q. clade, andcritically they did not sample type generaof many higher taxa. Given that themonophyly of many named groups
remains questionable, it is crucial tosample the type genera of each family,
subfamily and tribe to assess thetaxonomic limits of a given category.
Furthermore, previous molecular studies
have used only a small number ofmolecular markers, usually one to three
gene regions (Wahlberg & Wheat, 2008).
1.4 Outline of the thesis
In this thesis, I employed the methodsof molecular phylogenetics using eightmarkers for genomic DNA extractions of
Lepidoptera to study the evolution of
Noctuoidea and reconstruct a skeleton
phylogenetic hypothesis for its major
lineages. The first chapter gives acomprehensive overview of the higherlevel phylogeny and evolutionary affinities
of noctuoid moths, in an extensive
sampling strategy of the entire Noctuoidea(I). It reveals a new high-level
phylogenetic hypothesis comprising six
major, well-supported lineages that areinterpreted as families (I). The second
chapter aims to understand the higher-level
phylogeny and elucidate the evolutionary
history of the moth family Erebidae, themost controversial group among the newly
circumscribed families (II). Erebidae is amassive clade and includes a diverse
groups exhibiting a broad range of feeding
behaviors, including those that can beconsidered ‘piercers’ of fruits or otherhosts (skin-piercers: hematophagous) and‘tear feeders’ (lachryphagous) (III).
Within butterflies and moths, adulthematophagy is limited to species within
the vampire moth genus Calyptra

12Introduction

Ochsenheimer, which are placed within thesubfamily Calpinae, Erebidae. Paper III
focused on the subfamily Calpinae using
both morphological and molecular data toreconstruct ancestral feeding behaviors
within Calpinae as well as whether fruitpiercing
behavior
andassociatedmodifications of the tongue have evolved
independently in different groups ofErebidae (III). The fourth chapter seeks to
elucidate the deep divergences andevolutionary relationships of the major
lineages within the moth family Nolidae(IV). As a result of expanded sampling, a
new lineage (i.e., Diphtherinae) within
Noctuoidea—which includes taxa ofpreviously
uncertain
affinity—wasrecovered. Diphtherinae is considered to be
the plesiomorphic sister lineage to the restof Nolidae (IV), thus a new phylogenetic
hypothesis for Nolidae is presented (IV).
The fifth chapter elucidates theevolutionary relationships of the basal

lineages of the moth family Noctuidae (V).
A summary flow chart is presented in Fig.
2, representing my thesis process with theirmain outputs.

Figure 2 A flow chart summarizing five chapters
included in this thesis.

Material and Methods

13

2. MATERIAL AND METHODS
2.1 Sampling strategy
The sampling strategy that I adopted
has treated the world fauna. I attempted toinclude representatives of all known major
lineages of quadrifine Noctuoidea fromevery biogeographic region. This strategy
is being used to establish priorities forongoing studies, to test further therobustness of the major clades ofNoctuoidea, both in relation to each otherand internally.
Based on the results of recent
publications (Fibiger & Hacker, 2005,
Lafontaine & Fibiger, 2006, Mitchell et al.,2006, Lafontaine & Schmidt, 2010), 152,
237, 35, 120 and 76 terminal taxa were
sampled as representatives of the mostrecognized Noctuoidea lineages for papers
I–V, respectively. Furthermore, there werenumerous unplaced taxa of uncertain statusthat were included in each paper. Indeed,
the allocation of the unplaced taxa into any
higher taxa of Noctuoidea, however
tentatively, could not be determined inprevious classifications. I was unable tosample/amplify a few rare taxa withrestricted distributions and/or low species
richness (e.g., Strepsimaninae, Afridinae,
Camptolominae). Where possible, a
representative of the type genus of eachlineage is included, but this was notpossible for a few tribes/subtribes, inwhich case a closely related genus was
selected.
Appendix 1 summarizes all 393
terminal taxa that are used in the five
papers with their voucher codes and
GenBank accession numbers.
To test the monophyly of the target
taxon under study in the different papers, I
included representatives from the mostclosely related taxa of other families of
Lepidoptera (papers I, II, IV, V) or
subfamilies of Erebidae (III). I rooted thecladograms, in different papers, withdifferent taxa, which represents what I
consider to be the putative sister family tothe remainder of the terminal taxa.
2.2 Molecular markers
The total genomic DNA from one or
two legs, dried or freshly preserved in 96%
ethanol, was extracted using the DNeasy
tissue extraction kit (Qiagen, Hilden,
Germany) following the manufacturer’s
instructions. For each specimen, I
sequenced cytochrome c oxidase subunit I(COI) from the mitochondrial genome, andelongation factor-1α (EF-1α), ribosomal
protein S5 (RpS5), carbamoylphosphatesynthase domain protein (CAD), cytosolic
malatedehydrogenase(MDH),
glyceraldehyde-3-phosphate
dehydrogenase(GAPDH),
isocitrate
dehydrogenase (IDH) and wingless genes
from the nuclear genome. For paper III, I
sequenced an additional gene, the D2
region of the nuclear ribosomal RNA(rRNA) 28S region. All genes—except
28S, which is multiple-copy and encodes
ribosomal RNA—are single-copy, proteincoding exons and have previously beenfound to be highly informative in
phylogenetic analyses of Lepidoptera at
higher taxonomic levels (Wahlberg et al.,2009, Mutanen et al., 2010, I, II).
DNA amplification (PCR) andsequencing protocols follow Wahlberg &Wheat (2008). Sequencing was performed
mainly with an ABI 3730XL capillary
sequencer (Macrogen, Seoul, Korea), and asmaller part with an ABI PRISMR
3130XL capillary sequencer (Turku,

14Material and Methods

Finland). The resulting chromatograms
were checked and DNA sequences alignedby eye using BioEdit v. 7.0.4.1 (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. To
minimize the risk of any kind of confusion
during the sequencing protocol and errors
in alignments, I constructed neighborjoining and Maximum Likelihood trees
separately for each gene region andchecked them carefully for identicalsequences and other doubtful patterns. In
addition, to minimize the risk of
misidentification, all the specimens werecross-checked with their DNA barcodes(COI) in BOLD (Barcode of Life Data
System, http://www.boldsystems.org/views/login.php) (Ratnasingham & Hebert,2007), where reference specimens wereavailable for many of the species used inthis study.
2.3 Phylogenetic analyses and character
optimizations
The gene regions were analyzed using
various phylogenetic approaches including
model-based (Maximum Likelihood, ML;
and Bayesian Inference, BI), and nonmodel-based (i.e., parsimony) methods.
Initially, the data matrices were analysedin various combinations using ML to
explore their phylogenetic signal. Single
genes were analyzed on their own, nuclear
genes were combined, third codon
positions were removed, data waspartitioned into mtDNA and nDNA andfinally the data was partitioned by generegions (8 partitions). The effects ofvarying taxon and gene combinations werecompared against the analyses of the full,combined and partitioned by gene data.
Based on these data explorations, it was
decided to include all genes and third

codon positions as well as to partition thedata by genes (8 partitions) in the ML
analyses, and by nDNA and mtDNA (twopartitions) in the BI analyses.
Parsimony analyses (MP) were
undertaken using New Technology
heuristic searches implemented in the
program, TNT v 1.1 (Goloboff et al.,2003).
New
technology
searches(Goloboff, 1999) consisted of Tree Fusion,
Ratchet, Tree Drifting and Sectorial
Searches
performed,
withdefault
parameters applied, until the minimal tree
was found 1000 times. All characters weretreated as unordered and equally weighted,
and robustness of the hypothesis wasassessed through the bootstrap (BP) with
1000 pseudoreplicates (Felsenstein, 1985).
In addition, in papers I and II clade
support was estimated by Bremer support(BS) (Bremer, 1988, 1994) using a script(Peña et al., 2006) in TNT. Model-based
phylogenetic analyses were performed
using ML and a GTR + Γ model was
selected as the most appropriate model ofsequence evolution for each gene partition
based on the Akaike Information Criterionusing FindModel (http://www.hiv.lanl.gov/
content/sequence/findmodel/findmodel.ht
ml). ML analyses were conducted using
the default settings on the web-server
RAxML III BlackBox (Stamatakis et al.,2008). ML bootstrap analysis with 1000
pseudoreplicates (Felsenstein, 1985) was
also conducted with RAxML III.
BI was not used for papers I and II. In
the other papers, BI analyses were carriedout using the software MrBayes v3.1
(Ronquist et al., 2005) on the freely
available Bioportal server (http://www.
bioportal.uio.no). The dataset was divided
into two partitions: mtDNA and nDNA, as
partitioning by gene resulted in poor

15

Material and Methods
mixing of chains and problems withconvergence of likelihoods. I modeled theevolution of sequences according to the
GTR + Γ model independently for the twopartitions using the “unlink” command in
MrBayes. The Bayesian analyses wereseparately run two times for five, 23 and20 million generations for papers III−V,
respectively, with every 1000th generation
sampled. Clade robustness was estimatedby posterior probabilities (PP) in MrBayes.
Convergence was determined when thestandard deviation of split frequencies
went below 0.05 and the PSRF (Potential
Scale Reduction Factor) approached 1, andboth runs had properly converged to astationary distribution after the burn-in

stage (which
generations).

was

1,000

sampledTo understand character evolution inhigher noctuoids, a character optimization
analysis based on parsimony was
undertaken in paper IV using the software
Mesquite v2.75 (Maddison & Maddison,2011). Ancestral state reconstructions andcharacter transformations were optimized
onto the topology resulting from theBayesian analysis (IV).
All
laboratory
procedures
andphylogenetic data analyses are detailed in
the original papers.

16

Results and Discussion

3. RESULTS AND DISCUSSION
3.1 Phylogenetic hypothesis for
Noctuoidea
In the initial study, I aimed, as a firststep, to reconstruct a robust phylogenetic
hypothesis for higher taxa of Noctuoidea.
The results strongly supported themonophyly of Noctuoidea. The majorgroups within the Noctuoidea clade all
sharedaparticularmorphologicalsynapomorphy—a metathoracic tympanalorgan. I found six strongly supported
major lineages within Noctuoidea thatdeserved family status. These are
Oenosandridae, Notodontidae, Euteliidae,
Erebidae, Nolidae and Noctuidae (Fig. 3).
The first two major lineages are wellrecognized taxa that have often been
considered families within Noctuoidea.
Oenosandridae are a small family, only
known from Australia, comprising eightspecies in four genera (Nielsen et al.,1996), which mainly feed on Myrtaceae(Miller, 1991). Notodontidae containapproximately 3,800 species (Nieukerken
et al., 2011) and occur worldwide. The
other four lineages have been split into as
many as 10 families, with arctiines,
lymantriines and nolines frequently being
considered to be sufficiently distinct fromthe rest to warrant full family status. My
phylogenetic hypothesis (I) placed
previously recognized families—arctiines,
lymantriines, aganaines, herminiines andmicronoctuines—into
thestrongly
supported
Erebidaeclade.
WithinErebidae, relationships of only a few
lineages (e.g., Arctiinae, Aganainae andHerminiinae) were well supported (I). The
low support for some nodes within

Erebidae and Noctuidae probably stems
from
high
levelsofhomoplasy(particularly in the third codon position)and very sparse sampling.
At this stage, with well-supportedmonophyletic groups established within aphylogenetic framework, the question
arose of how best to apply the Linnaeansystem of nomenclature to the structure ofthat framework, by making decisions on
the content and arrangement of families,
subfamilies, tribes and subtribes in amanner that was most likely to optimize
the stability of that system and facilitate
access to its information content (II). In
particular, the establishment of a wellfoundedfamily-levelnoctuoidclassification is certainly an issue ofconsiderable practical importance, because
it affects the classification of about aquarter of the world’s lepidopteran species,
according to current estimates (Nieukerken
et al., 2011). In paper II, I discussed indetail reasons for applying the six familygroup system to the higher systematics ofNoctuoidea with their advantages anddrawbacks.
One of the most striking features of theapplication of molecular data in my thesis
was
uncoveringthephylogenetic
relationships of many unplaced taxa of
uncertain affinity. Molecular phylogenetic
techniques allowed the easy allocation ofthese taxa, which had remained ‘incertae
sedis’ or with uncertain limits for many
years. For example, the position of anumber of taxa characterized by thepseudoquadrifine hindwing venation had

Results and Discussion

17

been unstable for a long time (IV–V).
Most
of
these
groups(i.e.,
pseudoquadrifine
Noctuidae)
werepreviously assumed to be related to erebidsubfamilies (Fibiger & Lafontaine, 2005,
Lafontaine & Fibiger, 2006) (II) or evenconsidered as distinct families (e.g.,
Pantheidae) (Kitching & Rawlins, 1998)(V). The results of papers IV–V placed
them in a basal position within the family
Noctuidae with strong support.

and BI). In paper I, Euteliidae were sister
to Noctuidae in ML analyses (Fig. 3), andsister to the other three families together in
MP analyses. Similarly, in papers IV–V,
Euteliidae were placed as sister to the restof the quadrifid clade with moderately
good support in MP, ML and BI analyses(Figs 4–5). In paper I, Nolidae were sister
to Erebidae in ML analyses, but form atrichotomy with Erebidae and Noctuidae in
MP analyses.

3.2 Pattern of relationships amongmajor lineages of Noctuoidea

The short internal nodes, with little orno support for many basal divergences, inall quadrifid Noctuoidea lineages (I, II,
IV, V), suggest that these groups
diversified rapidly within a relatively shortperiod of time (Whitfield & Kjer, 2008).
Evolutionary rates in basal divergences
appear punctuated and such explosive
radiations are generally interpreted in two
different ways: (i) there is an historicalexplanation (e.g., rapid radiation or thesignature of a mass extinction event) or (ii)
it is simply an artefact. Under the rapid
radiation scenario, it is hypothesized thatthe lineages diverged so rapidly and withinsuch a narrow time window that there waslittle opportunity for the ancestors of eachmonophyletic group to evolve distinctive
apomorphies (Futuyma, 2005). However,such a pattern of short internal nodes andlineages with low support can be producedby other factors, such as inadequate data
quality, poor and sparse sampling strategy,
conflict within or among datasets (datainconsistency), incongruence between thereal evolutionary process and the assumed
models of sequence evolution, or even lackof phylogenetic signal due to accumulationof overlapping mutations (i.e., theprobability of substitutional saturation at agiven site).

Notodontidae are found to be the sistergroup of all other Noctuoidea, with
Oenosandridae branching off next (I).
However, this pattern of relationships
relative to the rest of Noctuoidea is notwell supported in all papers. BothOenosandridae and Notodontidae have atrifid forewing venation similar to that ofGeometridae, a character state that appears
to be plesiomorphic relative to thequadrifid forewing venation found in theother noctuoid families. All these terms are
discussed and defined in detail in paper II.
The results of paper I recovered the six
recognized families within Noctuoidea andthe monophyly of the quadrifid Noctuoidea(i.e., Euteliidae, Erebidae, Nolidae, andNoctuidae). Although, the relationships
amongst the remaining four families are
not clear, they formed a monophyletic
group (i.e., quadrifid Noctuoidea clade)with very strong support (Fig. 3) andshared a synapomorphy (i.e., quadrifid
forewing venation). The relationships ofthe four clades of quadrifid lineage
remained somewhat ambiguous in papers
I–II, although the results of papers IV–V
suggested a basal position for Euteliidae inall three phylogenetic methods (MP, ML18

Results and Discussion

Figure 3 The phylogenetic hypothesis of the superfamily Noctuoidea based on a maximum likelihood
analysis, along with outgroups. Clades representing families are coloured. The six families recognized here
are indicated. Names of moths shown in figure clockwise are: Notodontidae: Phalera Hübner; Euteliidae:
Eutelia Hübner, Noctuidae: Eucocytia Rothschild & Jordan (Pantheinae), Periphanes Hübner (Heliothinae);
Nolidae: Eligma Hübner (Eligminae); Erebidae: Scoliopteryx Germar (Scoliopteryginae), Lymantria Hübner
(Lymantriinae), Peridrome Walker (Aganainae), Euplagia Hübner (Arctiinae), Calyptra Ochsenheimer(Calpinae), Phytometra Haworth (Boletobiinae), Spirama Guenée (Erebinae), Cocytia Boisduval (Erebinae),
and Ophiusa Ochsenheimer (Erebinae).

In addition, a recent study indicates thatsuch phylogenetic patterns may well besignature of mass extinction events (Crisp& Cook, 2009), because mass extinction
produces a sharp drop in the cumulative
fossil diversity and is commonly thought tostimulate subsequent adaptive radiation,

creating a sharp increase in the rate ofdiversification (Benton & Emerson, 2007).

Results and Discussion

3.3 Phylogenetic hypothesis for
quadrifid Noctuoidea
Paper I revealed an urgent need for acomprehensive series of revisions for thehigherclassificationsofquadrifid
Noctuoidea families used up to that time.
My results in paper I almost failed torecover some previously recognizedsubfamilies
withinErebidaeasmonophyletic groups. For instance, theysuggested that some recent concepts of
subfamilies
Calpinae,
Catocalinae,
ErebinaeandPhytometrinae
werepolyphyletic (I). Consequently, the focus
in paper II was designed to elucidate thehigher-level phylogeny and evolutionary
relationships of the massive Erebidae clade(II). I thus conducted a large-scale
molecular phylogenetic analysis, which
uncovered a well-resolved skeletal
phylogenetic hypothesis. I thus presented a
new phylogenetic hypothesis for Erebidae
consisting of 18 moderate to strongly
supported
subfamilies
(Fig.
4):
Scoliopteryginae, Rivulinae, Anobinae,
Hypeninae, Lymantriinae, Pangraptinae,
Herminiinae,
Aganainae,
Arctiinae,
Calpinae, Hypocalinae, Eulepidotinae,
Toxocampinae,
Tinoliinae,
Scolecocampinae,
Hypenodinae,
Boletobiinae and Erebinae (Fig. 4). Where
possible,I
diagnosed
apomorphic
morphological character states for eachmonophyletic lineage (II).
Paper II provides strong support for
subordinating five taxa where previously
treatedasfamilies—Arctiidae,
Lymantriidae,
Micronoctuidae,
Herminiidae
andAganaidae—within
Erebidae. One of the most striking
features, a strong association among threeof them, was shown in papers I–II:
Aganainae + Herminiinae + Arctiinae are19

recovered as monophyletic clade (Fig. 4),
in which Arctiinae have a sister
relationship with a strongly supported
pairing of Aganainae and Herminiinae.
The clade also has a morphologicalsynapomorphy in the prespiracular positionof the counter-tympanal hood, which was
until then thought to be plesiomorphic.
Adults of many aganaines and arctiines arevisually striking and aposematic, andaganaines and herminiines share longlabial palps and a bare lower frons.
Herminiinae are generally cryptic, feedingon vegetable detritus, and Aganainae areaposematic, feeding on the same suite of
toxic
cardenolide-synthesizing
plantfamilies (Apocynaceae and Moraceae) as
the danaine Nymphalidae and other mothgenera such as Glyphodes (Crambidae) andAgathia (Geometridae) (Holloway, 2008).
Another interesting result from paper II
was the placement of the new establishedfamily Micronoctuidae. Hypenodinae areenlarged to incorporate Micronoctuini as atribe, corroborating the findings of paper I.
The subfamily Erebinae, brings together
the core catocalines (with the exceptions of
Hypocalinae,
ToxocampinaeandTinoliinae).
Calpinae sensu Lafontaine & Fibiger(2006) consisted of four tribes, Anomini,
Scoliopterygini, Calpini and Phyllodini on
the basis of the sharing of peculiar
morphological adaptations. A robust,
highly developed fruit-piercing (and insome cases skin-piercing and blood
sucking) proboscis is found widely in
Calpini and in some of the more robust
scoliopterygines such as Anomis Hübner,with many similar features in the structure.
However, my phylogenetic analysis

20

Results and Discussion

Figure 4 Phylogenetic hypothesis of the moths family Erebidae, based on ML analysis. Clades representing
the major clades are coloured. Support values (bootstrap) are shown next to the branches. Names of moths
shown in figure clockwise are: Nolidae: Eligma (Eligminae); Noctuidae: Periphanes (Heliothinae); Erebidae:
Scoliopteryx (Scoliopteryginae), Anoba Walker (Anobinae), Lymantria (Lymantriinae), Pangrapta Hübner
(Pangraptinae), Peridrome (Aganainae), Euplagia Hübner (Arctiinae), Syntomis Ochsenheimer (Arctiinae),
Calyptra (Calpinae), Eulepidotis Hübner (Eulepidotinae), Eublemma Hübner (Boletobiinae), Phytometra(Boletobiinae), Thysania Dalman (Erebinae), Catocala Schrank (Erebinae), Cocytia (Erebinae), and Ophiusa(Erebinae).

confirmed the polyphyly of the old conceptof Calpinae, and supports a monophyletic
Calpinae that places members of Anominiand Scoliopterygini in other noctuid
subfamilies (II–III). These results

suggesting that the fruit-piercing behavior
and the associated modifications of the
tongue seen in moths of both groups have
evolvedindependently(III).
The
phylogenetic hypothesis suggested threeResults and Discussion
subclades for the subfamily Calpinae thatwere treated as tribes: Phyllodini,
Ophiderini and Calpini (II). The polyphylyof the former concept of Calpinae provides
an object lesson in how the sharing of
peculiar morphological adaptations may
mislead classifications, and how sharedfeatures of a more subtle nature may beoverlooked in an unchallenged traditionalclassification.
WithintheentireLepidoptera, adult hematophagy—the
ability to pierce mammalian tissue andextract a blood meal—is limited to species
within the vampire moth genus Calyptra
Ochsenheimer, which belongs to thesubfamily Calpinae. In paper III, we tested
whether hematophagy in Calyptra arosefrom plant (e.g., fruit-piercing) or animalrelated behaviors (e.g., tear feeding or
lachryphagy). To do that, we subjected theresulting phylogenetic trees to a Bayesian
method of ancestral state reconstruction toreconstruct ancestral feeding behaviors
within Calpinae and test competing
hypotheses regarding their evolution. The
results supported the hypothesis that blood
feeding in vampire moth evolved from thefruit-piercing habit as opposed to tearfeeding or other animal-related feeding
behaviors (e.g., dung feeding, urine
feeding) (III).
In paper IV, I aimed to elucidate thehigher-level phylogeny of Nolidae and toclarify relationships in basal lineages ofNoctuidae(V).
My
phylogenetic
hypothesis (I) had already recovered
Nolidae and Noctuidae as well-supportedmonophyletic clades (Fig. 3). The results
were fascinating and fairly robust. Many
genera previously placed in Nolidae andtheformersubfamily
Ophiderinae(Erebidae) were placed with strong support
within Noctuidae, in the subfamily
Bagisarinae, supporting an expanded

21

concept of the subfamily (V). In addition,by increasing taxon sampling of several
unassigned Neotropical taxa, I uncovered apreviously unknown lineage of Noctuoidea
with Neotropical origins (Diphthera
Hübner + Lepidodes Guenée) with strong
support. This lineage appears to be thesister group to Nolidae, and it shares anunambiguous feature and the mostcharacteristic Nolidae apomorphy yet
proposed—the structure of the boat-shaped
cocoon with a vertical, anterior exit slit—,
suggesting that the clade could be includedin the family Nolidae as the subfamily
Diphtherinae (IV). Diphtherinae isconsidered to be the plesiomorphic sister
lineage to the rest of Nolidae,
characterized by the loss of the proximal
pair of tibial spurs on the hindlegs ofmales, and the presence of a frontal
tubercle or process, which is presumably
associated with a derived strategy ofemergence from the cocoon (IV). My
analyses (IV) revealed a well-resolved
phylogenetic hypothesis for Nolidae,
allowing me to present a new classification
for Nolidae consisting of eight strongly
supported
subfamilies:
Diphtherinae,
Risobinae,
Collomeninae,
Beaninae,
Eligminae, Westermanniinae, Nolinae, andChloephorinae. Among these, two groups
are suggested as new subfamilies (i.e.,
Collomeninae and Beaninae). I also
defined each monophyletic group by
autapomorphic morphological character
states.

3.4 Character optimizations
The higher systematics of the quadrifid
Noctuoidea is often discussed in terms ofwhether the hindwings have a trifine or
quadrifine venation. To understandcharacter evolution in higher noctuoids andmajor nolids, I undertook a character
optimization analysis based on parsimony

22

Results and Discussion

using the software Mesquite v2.75(Maddison & Maddison, 2011). My resultsof ancestral-state reconstructions andcharacter optimization (Fig. 5) showed thatmost traits—those characters that areconsidered to be of phylogenetic
significance—are clearly synapomorphic
for major lineages of quadrifids, except thehindwing venation which was an
ambiguous character state (IV). It is
generally
thoughtthatthepseudoquadrifine condition is the ancestral
state for Noctuoidea. This condition
concerns the position of three veins (M2,
M3 and CuA1), in which M2 arises about
one-third of the way up the discal cell, andM2 is strong and parallel to M3. My
character examinations indicated that thepseudoquadrifine condition is shared by
the basal lineages of Noctuidae s.s. (i.e.,
Bagisarinae, Plusiinae, Dilobinae andPantheinae) (V), and a few erebidsubfamilies (Hypeninae, Herminiinae,
Scoliopteryginae and Rivulinae) (II), as
well as Diphtherinae (Nolidae) (IV). I
evaluated the distribution patterns andevolutionary trends of this complex trait
under the two most commonly usedcharacter optimization algorithms ofparsimony
analysis:
ACCTRAN—accelerated
transformation—and
DELTRAN—delayedtransformation—(Agnarsson & Miller, 2008). I finallyfavoured DELTRAN optimization (IV),which minimizes reversals and maximizes
convergences and parallel evolution. Thisfavours the acquisition of the derived
quadrifine state—base of M2 close toM3—independently in Erebidae, Nolidae(with Diphtherinae excluded), andEuteliidae (Fig. 5). In other words, theplesiomorphic condition of M2 in thehindwing of the four quadrifid lineages isthe form found in Diphtherinae and basal
lineages of Noctuidae, and arguably somebasal lineages of Erebidae. This trend ofcharacter evolution can be traced bychecking the position of the vein M2 indifferent groups of quadrifids. For
instance, the condition of M2 in
Diphtherinae is exactly the same condition
that occurs in Pantheinae, Plusiinae,
Dilobinae and Bagisarinae (V), where M2
is very slightly reduced. In higher noctuids(i.e., Hadeninae) the vein is still visible inexactly the same position, but is even more
reduced, and then entirely lost in
Noctuinae (Fibiger & Lafontaine, 2005). In
the Erebidae lineage, M2 is seen in thiscondition in some primitive lineages (e.g.,
Rivulinae, Hypeninae, and Herminiinae).
As a consequence, the quadrifine (M2adjacent to M3) form of venation musthave been independently gained several
times within Erebidae, acquired once in
Nolidae (after the Diphtherinae lineage
branched off), and once in Euteliidae (Fig.
5). Thus, all these so-called primitive
lineages have retained the ancestral(symplesiomorphic) hindwing venation
trait with a pseudoquadrifine condition(Fig. 5).

3.5 Evolution of host-plant
associations in Noctuoidea
There have been few attempts to study
the evolution of host-plant use in
Noctuoidea. The results from the character
optimizations (IV) drove me to learn more
about the evolution of feeding habits inhigher noctuoids. One feature that has beensuggested as the prime factor governing
the evolution of butterfly-host plant
associations, is host growth form, whichappears, on the whole, to be moreconserved phylogenetically than host-planttaxon affiliation (Janz & Nylin, 1998).
Given its prevalence among the deeper
lineages, woody-plant feeding can be

Results and Discussion

23

Fig. 5 Summary of ancestral states reconstruction on Bayesian tree under DELTRAN optimization. Clades
representing higher taxa (i.e., families) are coloured. Support values under the two support measures(bootstrap/posterior probabilities) shown next to the branches. Coloured characters on terminal branches andinternal nodes indicate the presence of morphological traits as shown below the tree. Names of moths shownin figure from top to bottom are: Spirama (Erebidae, Erebinae), Autographa Hübner (Noctuidae, Plusiinae),
Diphthera (Nolidae: Diphtherinae), Eligma Hübner (Nolidae: Eligminae), Nola Leach (Nolidae, Nolinae),
Pseudoips Hübner (Nolidae, Chloephorinae, Chloephorini), Giaura Walker (Nolidae, Chloephorinae,
Sarrothripini).

reasonably inferred as ancestral for entire
Noctuoidea (Holloway, 1989, Mitchell etal., 2006), as it is, apparently, for mostother macrolepidopteran(Powell et al., 1998).

superfamilies

24

Results and Discussion

Within Noctuoidea, arboreal feeding is
predominant among the oldest and basal
lineages in trifid families—Oenosandridae
and Notodontidae—and, within quadrifids,in diverse lineages of Erebidae (e.g., many
Erebinae, Aganainae, Lymantriinae, etc.)
as well as Euteliidae (Table 1). My results
corroborate those of previous results(Forbes, 1923, Holloway, 1987, Weller,
1989, Miller, 1991, Richards, (1933) 1932)
in accepting that trifids are theplesiomorphic sister lineage to the rest of
Noctuoidea (i.e. quadrifid Noctuoidea) (I,II, IV, V). For example, the family
Oenosandridae—only
known
from
Australia—mainly feeds on the woody
plant family Myrtaceae (Miller, 1991). In
Notodontidae—withaworldwide
distribution but more diverse in tropics andespecially the Neotropics (Weller, 1989,
Miller, 1991)—almost all species feed ontrees, and only a few are found on
herbaceous plants (Miller, 1991). Within
Euteliidae, Euteliinae most commonly feedon Anacardiaceae (Powell et al., 1998), a
plant family that contains trees and shrubswith highly poisonous flowers. Among
their
other
prominenthosts
areBurseraceae (includes both shrubs andtrees), Dipterocarpaceae (mainly tropicallowland rainforest trees), Moraceae andHamamelidaceae (which consists of small
trees and shrubs) (Holloway, 1985, Powell
et al., 1998). Stictopterinae, the sistergroup of Euteliinae, is associated primarily
with Dipterocarpaceae and Clusiaceae(Table 1). Within quadrifids, where tree
feeding is probably also ancestral, it seems
clear that there have been many
independent colonizations and subsequent
radiations on herbaceous plants, mostspectacularly in Arctiinae (Erebidae) and
derived trifine lineages (e.g., the pest
clade). Although, the great majority ofquadrifids feed on living higher plants,

consumption of lower plants and detritus
has arisen in several groups, most notably
Erebidae (II). For example, lichen-feedingispredominant
among
lithosiines(Arctiinae) and a number of Aventiini(Boletobiinae) species (Wagner et al.,2008), while detritivory, mycophagy andalgivory is dominant in Herminiinae and
Boletobiinae (II) and recurs sporadically inother noctuid subfamilies (e.g., some
Bryophilinae, see Table 1) (V).
The results in paper V also suggested
that Noctuidae included a number oflineages that are exclusively arboreal
feeders, such as Dilobinae (Rosaceae),
Raphiinae(Salicaceae),
Pantheinae(Pinaceae) and Acronictinae (polyphagous
but mostly on trees) (Table 1). The first
three of these groups are associated withbasal Noctuidae lineages in my
phylogenetic hypothesis (Fig. 6)—thosecharacterized by the plesiomorphic
pseudoquadrifine
conditionofthehindwing venation. There are also a few
groups among the derived noctuid lineages
that generally feed on trees and shrubs,
such as some Xylenini, Psaphidini(Mitchell et al., 2006), and Orthosiini.
Some other subfamilies are more
specialized; for example, Agaristinae,
often show a strong preference for
Vitaceae (Holloway, 1989). My review ofmajor trends in feeding habits in thesubfamily Dyopsinae (Table 1) indicatesthat most of them feed on the plant family
Urticaceae, a flowering plant family of
mostly trees and shrubs. In contrast,
herbaceous-feeding larvae are predominant
in the higher trifines (i.e., the ‘pest clade’,Amphipyrinae, Metoponiinae, etc.).

Results and Discussion

25

Table 1 Larval host-plant families for Noctuidae study taxa. Zoogeographic regions are abbreviated asfollows: P = Palaearctic; O = Oriental; Au = Australasia; Nea = Nearctic; Neo = Neotropical; Af =Afrotropical.

26
Continuation of Table 1.

Results and Discussion

Results and Discussion
These results are also widely
corroborated by the study of Mitchell et al.
(2006), who studied the role of ecologicalandgeographicalfactors
inthediversification of Noctuoidea. They
presented a provisional synopsis of species
diversity, latitudinal distribution, and hostplant use for major noctuoid groups
sampled in their study, superimposed onto
the phylogeny. It is revealed that thegrowth form (i.e., woody-plant feeding vs.
herb-feeding) of the host-plant appears on
the
whole
more
conserved
phylogenetically than host-plant taxon
affiliation, where woody-plant feeding canbe reasonably inferred as ancestral for
Noctuoidea, as it is, apparently, for mostother macrolepidopteran superfamilies.
In general, herbaceous-feeding larvae
are predominant in the higher trifines (i.e.,
the
‘pest
clade’,
Amphipyrinae,
Metoponiinae, etc.), whereas in the basal
noctuid and basal trifines lineages arboreal
feeding is predominant. This evolutionary
pattern of feeding habits, postulating ageneral evolutionary trend from a tree
feeding to a herb feeding habit, has also
been shown for butterflies (Janz & Nylin,1998). Thus it can be suggested that thehost-growth form in Noctuoidea is more

27

evolutionarily conservative than hostaffiliation (Mitchell et al., 2006).
However, my preliminary study of hostplant associations showed that in a few
Noctuidae subfamilies a strong preference
toward feeding on a specific plant family
can be seen, such as Bagisarinae which
feed mainly on Malvaceae.
One of the fundamental reasons forstudying the evolutionary history of
noctuoid moths is to determine the main
driving forces behind the diversification ofthis species-rich group. It has become
apparent that the evolution of host-plant
use has likely been a key ecologicalmechanism behind the rapid diversificationand evolutionary divergence in thebutterfly families (Janz & Nylin, 1998,
Janz et al., 2006). Noctuoidea are about
three times more diverse than thebutterflies, and the role of host plant
specialization in the diversification of themoths has yet to be studied in detail. With
the phylogenetic relationships of the major
lineages of Noctuoidea becoming clearer(I, II, IV, V), questions about host plant
associations and diversification can now be
addressed for this megadiverse clade.

Fig. 6 Phylogenetic hypothesis of the basal Noctuidae subfamilies, based on a Bayesian inference analysis.
Clades representing major lineages are coloured. Support values under the two support measures(Bootstrap/posterior probabilities) shown next to the branches. Names of moths shown in figure clockwise
are: Eutelia (Euteliidae); Catocala (Erebidae); Eligma (Nolidae); Dyops (Noctuidae Dyopsinae, Dyopsini),
Sosxetra Walker (Dyopsinae, Ceroctena clade), Diloba Boisduval (Dilobinae), Eucocytia (Pantheinae),
Amyna Guenée (Bagisarinae), Vespola Walker (Bagisarinae), Concana Walker (Bagisarinae), Acronicta
Ochsenheimer (Acronictinae), Periphanes (Heliothinae), Euxoa Hübner (Noctuinae).

28

Results and Discussion

Conclusions and future directions

29

4. CONCLUSIONS AND FUTURE DIRECTIONS
4.1 Conclusions
To conclude my Ph.D. thesis, I havesummarized a number of distinctive issues
and strategies that I have employed in thisthesis that are different from those inprevious works.
First of all and probably the mostimportant scheme was the taxon samplingstrategy. Elucidating the evolutionary
history of the massive superfamily
Noctuoidea clade (potentially including
45,000 species) required extensive taxon
sampling. A deliberate and carefully
considered sampling strategy could only be
accomplished by selecting exemplars fromsignificant groupings of genera andmorphologically well-supported conceptsof higher taxa. In my five projects, I choseup to 393 taxa (Appendix 1) of 45,000
noctuoids that have been formallydescribed, covering almost all recognized
major
cladesofNoctuoidea,
asrepresentatives for major lineages ofNoctuoidea. This extensive taxon sampling
was made possible through the extensive
network that I built up during my studies.
One of the main causes for the previous
low support of phylogenetic relationships
for some massive clades probably stems
from very sparse sampling.
The second feature was the obstacles
that are met in applying morphologicaltraits, in particular, in a species-rich,cosmopolitan and heterogeneous taxon
such as Noctuoidea. The high number of
species presents complications to the useof morphology and any other kinds ofphenotypic traits at this taxonomic level.
Morphological data are thus often difficult
to homologize and code, require great
experience to identify character statescorrectly and can be subject to extensive
homoplasy (character convergence andreversal). Consequently, despite theirmajor role in inferring phylogenies,
morphological analyses have often failed
to determine relationships among mostgroups with confidence. On the other hand,
phenotypic traits (e.g., morphological,
ecological,
host-plant
associations,behaviouralcharacters,
etc.)
and
synapomorphiescanbeproperly
recognized from molecular phylogenybased systems. In the context of such arobustevolutionary
hypothesis,
morphological, ecological and behaviouralcharacters can be better understood. For
example, I demonstrated that theprespiracular counter-tympanal hood of
Aganainae, Herminiinae and Arctiinae is
not the result of convergent (independent)
evolutionary events, as previously thought(I–II). I proved that this is the result of acommon ancestry of these groups within
the family Erebidae (II). Polyphyly of theold concept of Calpinae was another clear
case (II–III). My results recovered amonophyletic subfamily Calpinae that was
restricted to three monophyletic tribes (i.e.,
Phyllodini, Ophiderini and Calpini) (III),and placed all other groups that had
previously been placed as tribes within
Calpinae (e.g., Anobini, Anomini and
Scoliopterygini), as independent anddistant
well-defined
lineages(i.e.,
Scoliopteryginae and Anobinae) within
Erebidae (II). Calpinae sensu lato was
traditionally restricted to the fruit-piercing(and in some cases skin piercing and blood
sucking) moths (Kitching & Rawlins,1998). However, my molecular phylogeny
revealed that this feeding behavior, as wellas its associated modifications of theproboscis and adaptations (III), have

30

Conclusions and future directions

evolved independently in Calpinae,
Scoliopteryginae (including Anomini),
Anobinae and some Erebinae (II). This
provides again an object lesson on how the
sharingofpeculiarmorphological
adaptations may mislead in classification,and how shared features of a more subtle
nature may be overlooked in an
unchallenged traditional classification (I).
Other
examplesofdiscoveringmorphological apomorphies based on mymolecular phylogeny results were in
Diphtherinae—characterized by the loss ofthe proximal pair of tibial spurs on thehindlegs of males and the presence of afrontaltubercle—(IV),
Nolidae—construction of a boat-shaped cocoon witha vertical exit slit, and finger-like
retinaculum on the forewings of themales—(IV),trifidandquadrifid
lineages—condition of vein M2 inforewing—(I, II, IV, V), host-plant
associations—e.g., detritivory, lichenfeeding, mycophagy and algivory—in
Boletobiinae (Erebidae) (II) and finally
speculation for a potential broad
evolutionary feeding habit trend in
Noctuoidea from tree feeding in trifid
lineages toward herb feeding in morederived lineages (i.e., pest clade in
Noctuidae), similar to the pattern that has
been suggested for butterflies (Janz &Nylin, 1998).
A third issue was related to determining
the evolutionary relationships of noctuoid
taxa of uncertain systematic position using
molecular data. In particular, those taxa
characterized by the pseudoquadrifine
hindwing venation, which have beenpreviously assigned to various noctuoidgroups (IV–V). Molecular phylogenetics
methods illustrate how easy is it can be to
pinpoint their certain phylogenetic
position.

To conclude, I had been able to addressamassiveandlong-recalcitrant
phylogenetic problem, that of therelationships among and within major
lineages of the largest superfamily of
Lepidoptera,
Noctuoidea.
Thephylogenetic analyses that I employed
were well designed, and many relatively
deep nodes are strongly resolved andsubstantial progress has been made on the
backbone phylogeny of Noctuoidea. This
group of insects is of major economic andecological importance. It constitutes an
exemplar case in which molecularmethods, which seem to be highly
informative at this level, have been ofenormous help, in part because the sheer
size of the group has greatly impeded
progress via the morphological approach.
4.2 Future directions
This Ph.D. thesis addressed several
phylogenetic problems concerning theevolution of Noctuoidea, but there are still
many unanswered questions. For example,
it is crucial to determine where, when andhow the major groups of noctuoids
diverged and evolved. The common factorsthat influence the speciation process andidentifying possible reasons for thisremarkable and extraordinary diversity of
species among other herbivorous insects
are not yet understood. Likewise, thepatterns of diversification, the main driving
forces behind the diversification of thisspecies-richgroup,
andplausible
explanations for the differences indiversity among the various groups within
Noctuoidea have yet to be determined—for
instance, why does a family such as
Euteliidae contain about 500 species, butanother like Erebidae contains 25,000
species? It would be of great interest to
know whether major climatic changes andmass extinction events over geological

Conclusions and future directions
time scales have had a major impact on thediversification of Noctuoidea. And it
would be most interested to infer ascenario of the evolutionary history andbiogeography of noctuoids, based on all
available data (i.e., morphology, DNA
sequences, ecological data, geographicaldistributions,andgeologicalandpaleontological information).
Further studies are also needed to
identify the reasons and causes for the
short basal branches, i.e., whether there isa historical explanation behind them (e.g.,
rapid radiation), or whether it is simply anartefact of insufficient data.

31

The use of novel tools in DNA
sequencing technologies, such as NextGeneration Sequencing (NGS) methods
and relatively new field of phylogenomics,
might be able to address the state ofuncertainty in Noctuoidea diversificationand their rapid radiations. Phylogenomics
is useful for evolutionary studies, inparticularresolvingambiguous
phylogenies and for verifying relationships
created on the basis of a few gene regions(Hackett et al., 2008). Since the cost ofwhole genome sequencing is decreasing,
anticipation of total genome sequencesfrom the major lineages of interest, is nolonger a distant dream (Murphy et al.,2004).

32

Acknowledgements

5. ACKNOWLEDGEMENTS
Laboratory work for this project was
funded through a grant from the Academy ofFinland (grant no. 129811) and Kone
Foundation to my supervisor, Dr. Niklas
Wahlberg. My subsistence expenses were
funded by the CIMO (May 2008 to April2009), the Finnish Cultural Foundation(February 2010 to January 2011) and theAlfred Kordelin Foundation (January 2012 toJune 2012). I would like to take thisopportunity to say thank to these Foundations
for supporting my four years stay in Finland.
Next, I would like to express my deep
thankfulness to my supervisor, Dr. Niklas
Wahlberg. To be honest, this dissertation
would not have been possible without Niklas’s
support and consistent guidance since the firstday I started my laboratory work (in September2008) to the final parts, where I am typing
these words (April 2012). I have started frombelow zero when Niklas taught me how towork in Genetics laboratory! It was acompletely new field to me as a taxonomist!
Niklas, you have been so kind all times, very
patient, always positive and constantly
optimistic. Thank you for the trust andconfidence you had on me.
I would like to dedicate this thesis to one of
the most influential persons in my life, the lateMichael Fibiger, who was my first teacher in
lepidopterology, God bless you kind man.
I owe my deepest gratitude to my thesis’
advisory team, Dr. Donald Lafontaine, Dr. Ian
Kitching and Dr. Jeremy Holloway, who madethis thesis possible. I think a nice feature of thisthesis was the inclusion of three leading
authorities on noctuoid morphology andclassification in the world, and that they havebeen closely involved in the design andinterpretation of my molecular results. Don, I
would like to thank you for all the discussions(including coffee and other supplies you
brought every day) and practical supervision
during the period I was working at the
Canadian National Collection of Insects. Don, I
would like to say, thank you for yourconfidence in me. Ian and Jeremy, I would liketo thank you for your fruitful, brilliant andcomprehensive comments on the manuscripts.
I am pleased to acknowledge my coauthors, Dr. Lauri Kaila and Dr. Marko
Mutanen, who have greatly contributed to theimprovement of manuscripts and also forsequencing of some material. I sincerely thankto Dr. Jennifer Zaspel, lead author of paper III,for helping me on that paper. I am grateful to
Dr. Chris Schmidt for all his practical andvaluable comments on our joint-papers (IV,V).
I also wish to acknowledge the work of Ms
Jocelyn Gill (CNC) for her expert work on thecolor plates for paper IV.
A large number of specimens for themolecular study were provided by a group of
nice people collaborating with us in thisproject: Prof Charles Mitter et al. (LepTree
project, University of Maryland, USA), Prof
Daniel H. Janzen (University of Pennsylvania,
USA); Roger C. Kendrick (Kadoorie Farm,
Hong Kong); Ugo Dall’Asta (Royal Museum
for Central Africa, Belgium); Lauri Kaila andJaakko Kullberg (Finnish Museum of Natural
History); Rob de Vos (Zoologisch Museum,
Netherlands); Erik Nieukerken (Netherlands
Centre for Biodiversity); Laszlo Ronkay(Hungarian Natural History Museum); Alexej
Matov (Zoological Institute of the Russian
Academy of Sciences); Leif Aarvik (Natural
History Museum, Norway); Shen-Horn Yen(National Sun Yat-sen University, Taiwan);
Henry Barlow (International Trust for
Zoological Nomenclature, UK & Malaysia);
Chris Muller (Flinders University, Australia);

Acknowledgements
Ian Kitching & Jeremy Holloway (Natural
History Museum, UK); Michael G. Pogue(Smithsonian Institution, U.S.A); Jennifer M.
Zaspel (University of Wisconsin Oshkosh,USA); Jérôme Barbut (Muséum nationald'Histoire
naturelle,
France);DonaldLafontaine and Christian Schmidt (theCanadian National Collection of Insects);
David Wagner (University of Connecticut,
USA); Vasiliy Kravchenko (Tel Aviv
University, Israel), as well as a number ofprivate collectors: Kari Nupponen (Espoo,
Finland); Petri Hirvonen (Porvoo, Finland);Szabolcs Safian (Hungary); Michael Fibiger(Sorø,
Denmark);
Jorg-Uwe
Meineke(Germany); Peter Smetacek (India); Ulf
Drechsel (Paraguay); and Markku Pellinen(Finland).
I wish to thank the pre-reviewers of thethesis, Prof. Charlie Mitter and Dr. Tommi
Nyman, for their informative comments on thethesis introduction. Despite not having had thechance to meet Prof. Mitter in person, I havelearned a lot from his great personality. I was
impressed, for the first time in October 2007,
before coming to Finland, when we wereasking experts to set up a strong research
proposal for PhD project. It is a pleasure to
thank Prof. Harri Savilahti and Dr. Erik van
Nieukerken, the custos and the opponent in mydoctoral disputation.
I would like to show my gratitude to Prof.
Craig Primmer for all his unseen but valuable
support. I am grateful to Ville Aukee and Meri
Lindqvist for all your assistance and technicalsupport.
My sincere thank goes to Dr. Carlos Peña,
NSG’ web application developer, who createdthe noctuid database for my thesis project.
Special thanks to my nice ex-officemates,
Akarapong Swatdipong (Pop) and Kalle
Rytkönen. Pop, thank you for all those nicemoments we have shared together in the sameoffice and coffee room. Kalle, thank you somuch for being positive, cheerful, joyful and33

helpful all the time. I will never forget ourFinnish sauna events, will never forget Vappu
2010 and 2011! I would like to thank Julien
Leneveu and Heike Witthauer for providing mewith the great atmosphere in which to work in
the TEGlab environment. I also wish to thankRaija Rouhiainen, who was the one who tookcare of my financial matters, calculating
Niklas’s grants several times a year.
I would like to thank Heidi and Johanna forkeeping the laboratory working. I owe sincere
thankfulness to Irma Saloniemi for all her
consultation, guidance and advice throughout
my doctoral studies. I would like to thank thelecturers and researchers of the ‘Laboratory of
Genetics’: Christina Nokkola, Seppo Nokkala,
Erica Leder, Anti Vasemägi, Juha-Pekka Vähä,
Sanna Huttunen, Mikko Nieminen, Spiros,
Matthieu, Olaf, and Susan. I am very gratefulto all the TEGlab people, in particular:
Tatjaana, Mikhail, Roghelio, Paula, Veronika,
Elsi, Heidi, Megha, Ksenia, Walter, Bineet andEero for technical help, resolving practicalissues, and of course tolerating me for four
years! Also, it is my privilege to share myfeelings of gratitude with my always cheerfulfriends, Pavel Matos and Siim Kahar. I would
like to express my sincere thanks to the UTU
Zoological Museum, especially Prof. Pekka
Niemelä, Ilari Sääksjärvi, Anssi Teräs (ÅA),
for your continued supports.
I am obliged to my wife, Maryam, who has
supported me through every situation. I also
want to thank my Aunt Shari for her support,
especially when I was deeply hopeless anddesperate.
And finally thanks to Suomi for letting me
be here for four years and allowing me finallyto live! I have experienced things I have neverexperienced before: ice swimming, enjoying 30
degrees below zero, smoked saunas, real
Finnish Saunas, Salmiakki vodka shots, getting
almost five months snow per year (Nov.-March
2010), spring snow, winter snow, autumn
snow!!!

34

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I
I

Bombyx moriArchiearis parthenias

Bombycinae

Archiearinae

Bombycidae

Geometridae

I,IV
I,II
I,II,IV
I,II,IV,V
I
I
I,V
I
I
I
I,II,V
IV
I
I,II,IV,V
II
I,II
IV,V
IV,V
I,II,IV
II
II
I,II
I,II
II
II
II
I,II

Phalera bucephala

Stauropus fagi

Notodonta dromedarius

Clostera pigra

Epicoma melanosticta

Thaumetopoea solitaria

Crinodes besckei

Nystalea striata

Scotura leucophleps

Eutelia adulatrix

Eutelia geyeri

Marathyssa basalis

Targalla subocellata

Aegilia describens

Lophoptera hemithyris

Lophoptera squammigera

Stictoptera columba

Scoliopteryx libatrix

Ossonoba torpida

Rusicada fulvida

Rusicada metaxantha

Gonitis involuta

Anomis flava

Rhesala imparata

Nychioptera noctuidalis

Rivula sericealis

Phalerinae

Heterocampinae

Notodontinae

Pygaerinae

Thaumetopoeinae

Thaumetopoeinae

Dudusinae

Nystaleinae

Dioptinae

Euteliinae

Euteliinae

Euteliinae

Euteliinae

Stictopterinae

Stictopterinae

Stictopterinae

Stictopterinae

Scoliopteryginae

Scoliopteryginae

Scoliopteryginae

Scoliopteryginae

Scoliopteryginae

Scoliopteryginae

unassigned

unassigned

Rivulinae

Notodontidae

Notodontidae

Notodontidae

Notodontidae

Notodontidae

Notodontidae

Notodontidae

Notodontidae

Notodontidae

Euteliidae

Euteliidae

Euteliidae

Euteliidae

Euteliidae

Euteliidae

Euteliidae

Euteliidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

I,II,IV

Oenosandra boisduvali

Oenosandridae

Discophlebia sp.

Oenosandridae

Ingroup

I
I

Thyatira batis

Sphinx ligustri

Thyatirinae

Sphinginae

Paper

Drepanidae

Outgroup

Species

Sphingidae

Subfamily

FamilyMM00027

Specimen ID
MM01404

RZ283

RZ265

RZ100

RZ13

RZ55

RZ101

RZ411

MM00407

RZ541

RZ120

MM07614

RZ287

RZ35

RZ23

RZ508

MM00160

06-SRNP-22781

05-SRNP-4443

05-SRNP-57213

MM09888

MM07592MM01005

MM00998

MM00981MM00122

RZ403

MM07590

NW107-1

NW149-1

NW141-12

COI-LCOGU828664

JN401256

JN401255

JN401254

HQ006166

HQ006227

JN401253

JN401252

GU828641

X

X

GU828802

JN401234

HQ006210

HQ006183

X

GU828621

GU828532

GU828525

GU828527

GU828843

GU828792

GU828654

GU828653

GU828651

GU828607

HQ006217

GU828791

DQ018928

EU141360

EU141358

GU828580

COI-Jerry
GU828462

JN401138

JN401137

JN401136

HQ006963

HQ006930

JN401135

JN401134

GU828439

X

X

GU929772

JN401118

HQ006914

HQ006887

X

GU828419

GU828334−−

GU929807

GU929763

GU828452

GU828451

GU828449

GU828405

HQ006921

GU929762

DQ018928

EU141360

EU141358

GU828380

íĮ-Eegin

EF1
GU828995

JN401373

JN401372

JN401371

HQ006263

HQ006322

JN401370

JN401369

GU828975

X

X

GU829107

JN401352

HQ006306

HQ006279

X

GU828956

GU828878

GU828871

GU828873

GU829144

GU829099

GU828985

GU828984

GU828983

GU828941

HQ006313

GU829098

DQ018899

EU136667

EU136665

GU828919

íĮ-enGEF1
GU829278−

JN401483

JN401482

HQ006357

HQ006414

JN401481

JN401480

GU829260

X

X

GU829385−

HQ006397

HQ006374

X

GU829246

GU829179

GU829173

GU829175−

GU829378

GU829269

GU829268

GU829266

GU829235

HQ006404

GU829377

DQ018899

EU136667

EU136665

GU829212

Wingless−

JN400941

JN400940

JN400981

HQ006775

HQ006835

JN400983

JN400939

GU829532

X

X

GU829661

JN400927

HQ006818

HQ006791

X

GU829516

GU829439

GU829432

GU829434

GU829692

GU829652

GU829541

GU829540

GU829539

GU829502

HQ006825

GU829651

DQ018869

EU141241

EU141239

GU829481

GAPDH
GU829791−

JN401585

JN401584−−

JN401583

JN401582−−

X

GU829879

JN401566

HQ006473

HQ006455

X

GU829764

GU829721

GU829717−

GU829904

GU829872

GU829782

GU829781

GU829780−

HQ006480

GU829871

EU141485

EU141495

EU141494

GU829743

RpS5
−

−

−

−

HQ006682

HQ006739

JN401894

JN401893

GU830643

X

X

GU830759

JN401873

HQ006722

HQ006698

X

GU830629

GU830568

GU830561

GU830563

GU830791

GU830752

GU830653

GU830652

GU830650

GU830617

HQ006729

GU830751

EU141381

EU141393

EU141391

GU830597

MDH
GU830370

JN401792

JN401791−

HQ006592

HQ006647

JN401790−

GU830348

X

X

GU830501

JN401772

HQ006631

HQ006606

X

GU830330

GU830256

GU830249

GU830251

GU830534

GU830493

GU830360

GU830359

GU830357

GU830318

HQ006638

GU830492

EU141604

EU141617

EU141615

GU830293

CAD
GU828161−

JN401053

JN401052

HQ006963

HQ007016

JN401051

JN401050

GU828140−−

GU828274

JN401036

HQ007000

HQ006979−

GU828122

GU828044

GU828037

GU828039

GU828307

GU828267

GU828151

GU828150

GU828148

GU828108−

GU828266

EU141303

EU141315

EU141313

GU828083

IDH
GU830051−

JN401685

JN401684−

HQ006560

JN401683−

GU830028

X

X

GU830183−−

HQ006528

X

GU830010

GU829923

GU829916

GU829918

GU830223

GU830174

GU830041

GU830040

GU830038

GU829995

HQ006551

GU830173

EU141539

EU141552

EU141550

GU829969

TG/TS

Type status
TG/TS

TS

TS

TG

TG

TS

TG/TS

TG

TG

TS

TS

TG

TG/TS

TG

TG

TG/TS

TS

TG/TS

TG/TS

TG/TS

TG/TS

TG/TS

FINLAND

Locality
FINLANDUSA

HONG KONG

HONG KONGTANZANIATAIWANHONG KONG

MALAYSIAFINLAND

MALAYSIA

HONG KONG

AUSTRALIA

INDONESIAHONG KONG

USA

HONG KONG

GREECECOSTA RICA

COSTA RICA

COSTA RICAGREECE

AUSTRALIAFINLANDFINLANDFINLANDFINLAND

AUSTRALIA

AUSTRALIASWEDENUSA

FINLAND

Appendix
37

APPENDIX

I,II
II
II
II
I,II

II
II
II
II
II
II
I,II

II
I,II
I,II,IV,V
I,II
I,II
I,II
I,II
I,II
I,II
II
II
II
II
I,II
II
I,II
I,II
I,II
I,II
II
II
II

Oxycilla ondo
Bocula bifaria
Oglasa ansorgei
Alesua etialis
Anoba anguliplaga
Marcipa sp.
Marcipa sp.
Plecoptera major
Crithote prominens
Rema costimacula
Baniana strigata
Deinopa signiplena
Hypena proboscidalis
Hypena baltimoralis
Hypena laceratalis
Cultripalpa sp.
Colobochyla salicalis
Lymantria monacha
Leucoma salicis
Nygmia plana
Orgyia antiqua
Arctornis sp.
Pangrapta bicornuta
Pangrapta decoralis
Chrysograpta igneola
Hyposemansis singha
Gracilodes caffra
Episparis costistriga
Masca abactalis
Schistorhynx argentistriga
Polypogon strigilatus
Paracolax tristalis
Herminia tarsicrinalis
Simplicia sp.
Idia aemula
Lysimelia neleusalis
Nodaria verticalis

Rivulinae

Rivulinae

Rivulinae

Rivulinae

Anobinae

Anobinae

Anobinae

Anobinae

Anobinae

Anobinae

Anobinae

Anobinae

Hypeninae

Hypeninae

Hypeninae

unassigned

unassigned

Lymantriinae

Lymantriinae

Lymantriinae

Lymantriinae

Lymantriinae

Pangraptinae

Pangraptinae

Pangraptinae

Pangraptinae

Pangraptinae

Pangraptinae

Pangraptinae

unassigned

Herminiinae

Herminiinae

Herminiinae

Herminiinae

Herminiinae

Herminiinae

Herminiinae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

II

II

I,II

II

Rivula ochrea

Rivulinae

Erebidae

RZ180

RZ260

RZ271

RZ166

RZ6

RZ5

MM01286

RZ119

RZ18

RZ319

RZ292

RZ279

RZ408

RZ66

RZ40

RZ89

RZ130

RZ34

MM06740

MM01048

RZ4

RZ394

RZ368

RZ367

MM01545

RZ311

RZ92

RZ103

RZ109

RZ183

RZ200

RZ177

RZ332

RZ94

RZ167

RZ413

RZ24

RZ159
JN401257

JN401277

JN401276

JN401275

HQ006175

HQ006232

HQ006224

GU828663

JN401274

HQ006178

JN401273

JN401272

JN401271

JN401270

HQ006236

HQ006216

HQ006241

HQ006167

HQ006209

GU828748

GU828655

HQ006215

JN401269

JN401268

JN401267

GU828668

JN401266

JN401265

JN401264

JN401263

JN401262

JN401261

HQ006177

HQ006206

JN401260

JN401259

JN401258

HQ006184

JN401139

JN401159

JN401158

JN401157

HQ006879

HQ006935

HQ006927

GU828461

JN401156

HQ006882

JN401155

JN401154

JN401153

JN401152

HQ006939

HQ006920

HQ006943

HQ006964

HQ006913

GU929722

GU828453

HQ006919

JN401151

JN401150

JN401149

GU828466

JN401148

JN401147

JN401146

JN401145

JN401144

JN401143

HQ006881

HQ006910

JN401142

JN401141

JN401140

HQ006888

JN401374

JN401392−

JN401391

HQ006272

HQ006327

HQ006319

GU828994

JN401390

HQ006274

JN401389

JN401388

JN401387

JN401386

HQ006331

HQ006312

HQ006335

HQ006264

HQ006305

GU829062

GU828986

HQ006311

JN401385

JN401384

JN401383

GU828999

JN401382

JN401381

JN401380

JN401379−

JN401378−

HQ006302

JN401377

JN401376

JN401375

HQ006280

JN401484

JN401504

JN401503

JN401502

HQ006366

HQ006419

HQ006411

GU829277

JN401501

HQ006369

JN401500

JN401499

JN401498

JN401497

HQ006423

HQ006403

HQ006428

HQ006358

HQ006396

GU829347

GU829270

HQ006402

JN401496

JN401495

JN401494

GU829282

JN401493

JN401492

JN401491

JN401490

JN401489

JN401488

HQ006368

HQ006395

JN401487

JN401486

JN401485

HQ006375

JN400979

JN400959

JN400958

JN400957−

HQ006840

HQ006832

GU829549

JN400956

HQ006786

JN400955

JN400954

JN400953

JN400952

HQ006844

HQ006824

HQ006849

HQ006776

HQ006817

GU829611

GU829542

HQ006823

JN400951

JN400950

JN400993

GU829553

JN400949

JN400948

JN400947

JN400946

JN400945

JN400944

HQ006785

HQ006814

JN400943

JN400986

JN400942

HQ006792

JN401586

JN401604

JN401603−

HQ006448

HQ006489−

GU829790

JN401602

HQ006451

JN401601

JN401600

JN401599

JN401598−

HQ006479

HQ006494

HQ006443

HQ006472−−

HQ006478

JN401597−

JN401596

GU829794

JN401595

JN401594

JN401593

JN401592

JN401591

JN401590

HQ006450

HQ006469

JN401589

JN401588

JN401587

HQ006456

−

−

−

−

HQ006691−

HQ006736

GU830660

JN401902

HQ006693

JN401901

JN401900

JN401899

JN401898

HQ006747

HQ006728

HQ006752

HQ006683

HQ006721

GU830719

GU830654

HQ006727

JN401897

JN401896

JN401895

GU830664

−

−

−

−

−

−

−

−

−

−

−

−

JN401793−

JN401808

JN401807

HQ006599−−

GU830369

JN401806−−

JN401805

JN401804

JN401803−

HQ006637

HQ006659

HQ006593

HQ006630

GU830449

GU830361

HQ006636

JN401802−

JN401801

GU830374

JN401800

JN401799

JN401798

JN401797

JN401796

JN401795

HQ006601

HQ006627

JN401794−−

HQ006607

JN401054−

JN401067

JN401066

HQ006971−

HQ007013

GU828160

JN401065

HQ006974−

JN401064

JN401063−

HQ007022

HQ007006

HQ007024

HQ006964

HQ006999

GU828232

GU828152

HQ007005−

JN401062

JN401061

GU828165

JN401060

JN401059

JN401058

JN401057−−

HQ006973−−

JN401056

JN401055

HQ006980−

JN401704

JN401703

JN401702

HQ006520−−

GU830050

JN401701

HQ006523

JN401700

JN401699

JN401698−

HQ006568

HQ006550

HQ006572

HQ006513

HQ006546

GU830132

GU830042−

JN401697

JN401696

JN401695

GU830055

JN401694

JN401693

JN401692

JN401691

JN401690

JN401689

HQ006522

HQ006544

JN401688

JN401687

JN401686

HQ006529

TS

TS

TG

TG/TS

TS

TS

TS

TS

TS

TG/TS

TG

TG

TG/TS

TG

TG/TS

TG

TS

TG

TG

TG/TS

TS

TG

TS

TG

GHANA

HONG KONG

USA

GHANA

HUNGARYHUNGARY

FINLAND

HONG KONG

INDONESIA

MALAYSIATANZANIA

MALAYSIA

MALAYSIAUSA

HONG KONGJAPANFINLAND

HONG KONG

FINLANDFINLANDHUNGARY

MALAYSIA

HONG KONG

USA

FINLANDCOSTA RICA

COSTA RICA

HONG KONG

HONG KONG

GHANA

GHANA

GHANA

GHANA

COSTA RICA

GHANA

MALAYSIA

USA

GHANA

38
Appendix

I,II
II
II
II
II
II
I,II
II
II

II
I,II
I,II
I,II
I,II
I,II
I,II
I,II
I,II
II
I,II
I,II
I,II,IV,V
II
II
I,II,III
II,III
I,II,III
II,III
II,III
I,II,III
III
III
I,II,III
III
III
III

Asota heliconia
Neochera inops
Euplocia membliaria
Peridrome orbicularis
Mecodina praecipua
Psimada quadripennis
Brunia antica
Garudinia simulana
Eugoa bipunctata
Cyana sp
Barsine sp
Apisa canescens
Syntomis phegea
Dysauxes famula
Antichloris viridis
Ctenucha virginica
Coscinia cribraria
Pseudophaloe troetschi
Dysschema leucophaea
Nyctemera baulus
Callimorpha dominula
Creatonotos transiens
Arctia caja
Amerila astreus
Epitausa dilina
Phyllodes eyndhovii
Phyllodes imperialis
Miniodes phaeosoma
Hemiceratoides sittaca
Eudocima salaminia
Eudocima fullonia
Eudocima divitiosa
Eudocima tyrannus
Gonodonta uxor
Gonodonta lincus
Gonodonta milla
Gonodonta syrna

Aganainae

Aganainae

Aganainae

Aganainae

Aganainae

Aganainae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Arctiinae

Unassigned

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

ErebidaeII

I,II

Asota caricae

Aganainae

Erebidae

RZ420

RZ421

RZ417

RZ335

RZ430

RZ210

RZ16

RZ338

RZ155

RZ153

RZ546

RZ56

RZ93

RZ404

MM03713

RZ30

RZ136

RZ387

RZ88

06-SRNP-35191

MM05671

AM-94-0396

MM05380

MM00154

RZ8

MM05843

RZ397

RZ398

RZ400

RZ399

RZ28

RZ47

RZ268

RZ280

RZ345

RZ346

RZ44

MM00145

JN674874

JN674873

JN674872

HQ006208

JN674871

JN674870

HQ006174

JN401291

JN401290

HQ006173

JN674869

HQ006228

JN401289

JN401288

GU828693

HQ006198

HQ006169

JN401287

HQ006240

GU828534

HQ006149

GU828535

HQ006151

GU828619

HQ006238

HQ006146

JN401286

JN401285

JN401284

JN401283

HQ006193

JN401282

JN401281

JN401280

JN401279

JN401278

HQ006220

GU828615

GU828413

JN674856

JN674855

JN674854

HQ006912

JN674853

JN674852

HQ006878

JN401173

JN401172

HQ006877

JN674851

HQ006931

JN401171

JN401170

GU828489

HQ006902

HQ006873

JN401169−

GU828336

HQ006856

GU828337

HQ006858

GU828417

HQ006941

HQ006853

JN401168

JN401167

JN401166

JN401165

HQ006897

JN401164

JN401163

JN401162

JN401161

JN401160

HQ006924

GU828949

JN674891

JN674890

JN674889

HQ006304

JN674888

JN674887

HQ006271

JN401406

JN401405

HQ006270

JN674886

HQ006323

JN401404

JN401403−

HQ006294

HQ006266

JN401402

HQ006334

GU828880

HQ006247

GU828881

HQ006249

GU828954−−

JN401401

JN401400

JN401399

JN401398

HQ006289

JN401397

JN401396

JN401395

JN401394

JN401393

HQ006316

GU829240

JN674907

JN674906

JN674905−

JN674904

JN674903

HQ006365

JN401517

JN401516

HQ006364

JN674902

HQ006415

JN401515

JN401514

GU829305

HQ006387

HQ006360

JN401513

HQ006427

GU829180

HQ006342

GU829181

HQ006344

GU829244

HQ006425

HQ006339

JN401512

JN401511

JN401510

JN401509

HQ006383

JN401508

JN401507

JN401506

JN401505−

HQ006407

GU829509

JN674973

JN674972

JN674971

HQ006816

JN674970

JN674969

HQ006783

JN400990

JN400971

HQ006782

JN674968

HQ006836

JN400970−

GU829573

HQ006806

HQ006778

JN400969

HQ006848

GU829441

HQ006758

GU829442

HQ006760

GU829514

HQ006846

HQ006765

JN400968

JN400967

JN400966

JN400965

HQ006801

JN400964

JN400963

JN400962

JN400961

JN400960

HQ006828−

JN674924

JN674923

JN674922

HQ006471

JN674921

JN674920

HQ006447

JN401615

JN401614

HQ006446

JN674919−

JN401613

JN401612

GU829813−

HQ006444

JN401611

HQ006493−−

GU829722

HQ006433−

HQ006492−−

JN401610−

JN401609

HQ006462−

JN401608

JN401607

JN401606

JN401605

HQ006483

GU830624

JN674955

JN674954

JN674953

HQ006720

JN674952

JN674951

HQ006690

JN401913

JN401912

HQ006689

JN674950

HQ006740

JN401911

JN401910−

HQ006711

HQ006685

JN401909

HQ006751

GU830569

HQ006666

GU830570

HQ006668−

HQ006749

HQ006663

JN401878

JN401876

JN401906

JN401905

HQ006706

JN401904−

JN401903−−

HQ006732

GU830325

JN674938

JN674937

JN674936

HQ006629

JN674935−

HQ006598

JN401821

JN401820

HQ006597

JN674934

HQ006648

JN401819−

GU830398

HQ006619

HQ006594

JN401818

HQ006658

GU830258−−

HQ006579

GU830328

HQ006656−

JN401817

JN401816

JN401815

JN401814

HQ006614

JN401813

JN401812

JN401811

JN401810

JN401809

HQ006641

GU828115

−

−

−

−

−

−

HQ006970−−

HQ006969

−

−

−

−

GU828185

HQ006991

HQ006965−−

GU828046

HQ006949−

HQ006951

GU828120−−

JN401074

JN401073

JN401072

JN401071−

JN401070

JN401069−−

JN401068

HQ007009

GU830003

JN674996

JN674995

JN674994

HQ006545

JN674993

JN674992

HQ006519

JN401740

JN401717

HQ006518

JN674991

HQ006561

JN401716

JN401715

GU830080

HQ006537

HQ006514

JN401714

HQ006571

GU829925

HQ006499

GU829926

HQ006501

GU830008−−

JN401713

JN401712

JN401711

JN401710

HQ006534

JN401709

JN401708

JN401707

JN401706

JN401705

HQ006554

TG

TG

TG

TG/TS

TG

TG

TS

TG/TS

TG/TS

TG

TG/TS

TS

TS

TG/TS

TS

TS

TG

TG/TS

TAHILANDBRAZIL

BRAZIL

BRAZILCOSTA RICARUSSIAGHANA

MALAYSIA

HONG KONG

GHANA

GHANA

AUSTRALIA

TAIWANCOSTA RICA

MALAYSIAFINLAND

HONG KONG

RUSSIAMALAYSIACOSTA RICA

COSTA RICA

FINLANDUSAECUADORGREECEHUNGARYOMAN

MALAYSIA

MALAYSIA

MALAYSIA

MALAYSIA

HONG KONG

HONG KONG

HONG KONG

MALAYSIA

MALAYSIAJAPAN

HONG KONG

Appendix39

III
I,II,III,IV,V MM00963

II,III
III
III
III
I,II,III
III
III
I,II,III
II,III
III
III
III
III
I,II,IIIII
II,IIIII
I,II,III
II,IIIII
I,II,III
II
II
II
I,II
I,II,III
II
I,II
I,II
II
II
II
II
I,II

Gonodonta sicheas
Calyptra thalictri
Calyptra hokkaida
Calyptra lata
Calyptra canadensis
Calyptra minuticornis
Plusiodonta nitissima
Plusiodonta coelonota
Plusiodonta casta
Oraesia emarginata
Oraesia excavata
Oraesia excavate
Oraesia nobilis
Oraesia glaucocheila
Oraesia rectristria
Hypsoropha hormos
Hypocala deflorata
Hypocala andremona
Tautobriga glaucopis
Panopoda rufimargo
Antiblemma fuscireticulata
Sanys irrosca
Eulepidotis rectimargo
Anticarsia gemmatalis
Anticarsia irrorata
Hemeroplanis finitima
Oxidercia toxea
Azeta ceramina
Autophila chamaephanes
Lygephila pastinum
Lygephila maxima
Tinolius eburneigutta
Poeta denotalis
Tamsia hieroglyphica
Platyjionia mediorufa
Scolecocampa liburna

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Calpinae

Hypocalinae

Hypocalinae

Hypocalinae

Eulepidotinae

Eulepidotinae

Eulepidotinae

Eulepidotinae

Eulepidotinae

Eulepidotinae

Eulepidotinae

Eulepidotinae

Eulepidotinae

Eulepidotinae

Toxocampinae

Toxocampinae

Toxocampinae

Tinoliinae

Tinoliinae

Tinoliinae

Unassigned

Scolecocampinae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

RZ9

RZ111

RZ389

RZ445

RZ331

RZ57

MM05092

RZ276

RZ22

RZ295

RZ298

RZ370

RZ267

RZ12

RZ343

RZ334

RZ59

RZ354

RZ340

RZ105

RZ17

RZ433

RZ418

RZ422

RZ434

RZ337

RZ102

RZ429

RZ106

RZ333

RZ514

CTW2

RZ431

RZ336

RZ419

RZ432

III

Gonodonta nutrix

Calpinae

Erebidae

RZ423

III

Gonodonta fulvangula

Calpinae

Erebidae

HQ006242

JN401306

JN401305

JN401304

JN401303

HQ006229

GU828711

JN401302

HQ006182

HQ006196

JN401301

JN401300

JN401299

HQ006162

JN401298

JN401297

HQ006231

JN401296

JN401295

JN401294

HQ006176

JN674883

JN674882

JN674881

JN674880

JN401293

HQ006159

JN674879

JN674878

HQ006207

JN674885−

JN674884

JN401292

HQ006156

JN674877

JN674876

JN674875

HQ006944

JN401188

JN401187

JN401186

JN401185

HQ006932

GU828506

JN401184

HQ006886

HQ006900

JN401183

JN401182

JN401181

HQ006960

JN401180

JN401179

HQ006934

JN401178

JN401177

JN401176

HQ006880

JN674865

JN674864

JN674863

JN674862

JN401175

HQ006864

JN674861

JN674860

HQ006911

JN674868

JN674867

JN674866

JN401174

HQ006861

JN674859

JN674858

JN674857

HQ006336

JN401420

JN401419

JN401418

JN401417

HQ006324−

JN401416

HQ006278

HQ006292

JN401415

JN401414

JN401413

HQ006259

JN401412

JN401411

HQ006326−

JN401410

JN401409

HQ006273

JN674899

JN674898

JN674897−

JN401408

HQ006256

JN674896

JN674895

HQ006303

JN674901−

JN674900

JN401407

HQ006252

JN674894

JN674893

JN674892

HQ006429

JN401528

JN401527

JN401526−

HQ006416

GU829323

JN401525

HQ006373−

JN401524

JN401523

JN401522

HQ006354−−

HQ006418−

JN401521

JN401520

HQ006367

JN674916

JN674915

JN674914

JN674913

JN401519

HQ006351

JN674912

JN674911−

JN674918−

JN674917

JN401518

HQ006348

JN674910

JN674909

JN674908

HQ006850

JN400991

JN401007

JN401006−

HQ006837

GU829587

JN401005

HQ006790

HQ006804

JN401004

JN400995

JN400974

HQ006771

JN400973

JN400975

HQ006839

JN400977

JN400980

JN400985

HQ006784

JN674981

JN674980

JN674979−

JN400987

HQ006768

JN674978JN674977

HQ006815

JN674984

JN674983

JN674982

JN400972

HQ006763

JN674976

JN674975

JN674974

HQ006495

JN401628

JN401627

JN401626−

HQ006487−

JN401625−−

JN401624

JN401623

JN401622−

JN401621

JN401620

HQ006488−

JN401619

JN401618

HQ006449

JN674930

JN674929

JN674928−

JN401617

HQ006439

JN674927

JN674926

HQ006470

JN674932

JN674933

JN674931

JN401616

HQ006435

JN674925−−

HQ006753

JN401926

JN401907

JN401925

JN401924

HQ006741

GU830699

JN401923

HQ006697

HQ006709

JN401922

JN401921

JN401920

HQ006678

JN401919

JN401918

HQ006743−

JN401917

JN401916

HQ006692

JN674964

JN674963

JN674962

JN674958

JN401915

HQ006675

JN674961

JN674960

HQ006719

JN674966

JN674967

JN674965

JN401914

HQ006671

JN674959

JN674956

JN674957

HQ006660−

JN401835−

JN401834

HQ006649

GU830415

JN401833

HQ006605

HQ006617

JN401832

JN401831

JN401830

HQ006588

JN401829

JN401828

HQ006651

JN401827

JN401826

JN401825

HQ006600

JN674946

JN674945

JN674944−

JN401824

HQ006586

JN674943

JN674942

HQ006628

JN674948

JN674949

JN674947

JN401823

HQ006582

JN674941

JN674939

JN674940−

HQ007025

JN401084

JN401083−

JN401082−

GU828199

JN401081

HQ006978

HQ006989−

JN401080−

HQ006960

JN401079−

HQ007018−

JN401078

JN401077

HQ006972−

JN674987−

JN674986

JN401076

HQ006958−−−

JN674990

JN674989

JN674988

JN401075

HQ006955−

JN674985

HQ006573

JN401729

JN401728

JN401727−

HQ006562

GU830097

JN401726

HQ006527−

JN401725

JN401724

JN401723

HQ006511

JN401722−

HQ006564−

JN401721

JN401720

HQ006521

JN675004

JN675003

JN675002−

JN401719

HQ006508

JN675001

JN675000−

JN675006

JN675005−

JN401718

HQ006504

JN674999

JN674998

JN674997

TG

TS

TG/TS

TG

TG

TS

TS

TG

TG

TG

TG/TS

TS

TG

TG

TG

TG

TG/TS

USA

HONG KONG

MALAYSIA

MALAYSIATHAILANDJAPAN

FINLANDCOSTA RICA

COSTA RICA

COSTA RICAUSA

HONG KONGUSACOSTA RICA

COSTA RICA

COSTA RICAUSA

COSTA RICA

COSTA RICA

HONG KONG

USA

NEPAL

BRAZIL

BRAZILUSA

HONG KONG

HONG KONG

RUSSIA

HONG KONGCOSTA RICA

MALAYSIAUSA

RUSSIAJAPANFINLANDECUADORUSA

BRAZIL

40
Appendix

I,II
I,II
II
II

II
II
I,II
II
I,II
I,II
I,II
II
II
II
II
II

II
I,II
II
II

Hypenodes humidalis
Schrankia costaestrigalis
Luceria striata
Luceria oculalis
Anachrostis sp.
Micronoctua sp.
Biuncus sp. 1
Biuncus sp. 2
Saroba pustulifera
Condate sp.
Corgatha nitens
Phytometra viridaria
Laspeyria flexula
Zurobata rorata
Homodes crocea
Enispodes purpurea
Tamba mnionomera
Parolulis absimilis
Araeopteron sp.
Araeopteron sp.
Eublemma purpurina
Eublemma anachoresis

Hypenodinae

Hypenodinae

Hypenodinae

Hypenodinae

Hypenodinae

Hypenodinae

Hypenodinae

Hypenodinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

I,II
II
I,II
I,II
II
I,II
II
II
I,II
I,II
II
I,II
II

Eublemma albifascia
Parascotia fuliginaria
Metalectra edilis
Trisateles emortualis
Prolophota trigonifera
Hypenagonia ?brachypalpia
Metaemene atrigutta
Mataeomera semialba
Euaontia semirufa
Acantholipes circumdata
Acantholipes regularis
Hypospila bolinoides
Ugia insuspecta
Ugiodes cinerea

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Boletobiinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

I,II

I,II

II

II

Gabara stygialis

Scolecocampinae

Erebidae

RZ326

RZ45

RZ116

RZ135

RZ248

RZ285

RZ107

RZ41

RZ409

RZ37

MM04877

RZ372

MM00340

RZ220

RZ98

RZ7

RZ410

RZ137

RZ392

RZ415

RZ390

RZ412

RZ385

RZ3

RZ129

RZ36

RZ393

RZ104

RZ476

RZ475

RZ138

RZ288

RZ369

RZ42

RZ27

MM01780

RZ297
JN401307

JN401326

HQ006221

JN401325

HQ006168

HQ006189

JN401324

JN401323

HQ006218

JN401322

HQ006212

GU828707

JN401321

HQ006154

JN401320

JN401319

HQ006237

JN401318

HQ006170

JN401317

JN401316

JN401315

JN401314

JN401313

HQ006197

HQ006165

HQ006211

JN401312

HQ006160

JN401311−

HQ006171

JN401310

JN401309

JN401308

HQ006192

GU828671

JN401189

JN401208

HQ006925

JN401207

HQ006872

HQ006893

JN401206

JN401205

HQ006922−

HQ006916

GU828502

JN401204

HQ006862

JN401203

JN401202

HQ006940

JN401201

HQ006874

JN401200

JN401199

JN401198

JN401197

JN401196

HQ006901

HQ006962

HQ006915

JN401195

HQ006865

JN401194

JN401193

HQ006875

JN401192

JN401191

JN401190

HQ006896

GU828469−−

JN401440

HQ006265

HQ006285

JN401439

JN401438

HQ006314

JN401437

HQ006308

GU829030

JN401436

HQ006253

JN401435

JN401434

HQ006332

JN401433

HQ006267

JN401432

JN401431

JN401430

JN401429

JN401428

HQ006293

HQ006262

HQ006307

JN401427

HQ006257

JN401426

JN401425

HQ006268

JN401424

JN401423

JN401422

HQ006288−

JN401421

JN401529−

HQ006408

JN401544

HQ006359

HQ006379

JN401543−

HQ006405

JN401542

HQ006399

GU829319

JN401541

HQ006347

JN401540

JN401539

HQ006424

JN401538

HQ006361

JN401537

JN401536−

JN401535

JN401534

HQ006386

HQ006356

HQ006398

JN401533

HQ006352−

JN401532

HQ006362−

JN401531

JN401530

HQ006382

GU829285−−

HQ006829

JN400997

HQ006777

HQ006797

JN401017−

HQ006826

JN401016

HQ006820

GU829583

JN401015

HQ006764

JN401014

JN400989

HQ006845

JN401013

HQ006779−−

JN401012

JN401011

JN400996

HQ006805

HQ006774

HQ006819

JN401010

HQ006769

JN400994

JN400992

HQ006780−

JN401009

JN401008

HQ006800

GU829556

JN401644

HQ006484

JN401643−−

JN401642

JN401641

HQ006481

JN401640

HQ006475

GU829821

JN401639

HQ006436

JN401638

JN401637

HQ006491

JN401636−

JN401635

JN401634

JN401633−−

HQ006463

HQ006442

HQ006474

JN401632−

JN401631

JN401630

HQ006445−−−

HQ006461−

JN401629

JN401927

JN401945

HQ006733

JN401944

HQ006684

HQ006702

JN401943−

HQ006730

JN401942

HQ006724

GU830695

JN401941

HQ006672

JN401940

JN401939

HQ006748

JN401938

HQ006686

JN401937

JN401936

JN401935

JN401934

JN401933

HQ006710

HQ006681

HQ006723

JN401932

HQ006676

JN401908

JN401931

HQ006687

JN401930

JN401929

JN401928

HQ006705

GU830666−

HQ006642

JN401851−−−

JN401850

HQ006639

JN401849

HQ006633

GU830411

JN401848

HQ006583−

JN401847

HQ006655

JN401846−

JN401845

JN401844

JN401843

JN401842

JN401841

HQ006618

HQ006591

HQ006632

JN401840−

JN401839

JN401838

HQ006595

JN401837−−

HQ006613−

JN401836

JN401085

JN401097

HQ007010−−

HQ006984

JN401096

JN401095

HQ007007−

HQ007002

GU828195−

HQ006954−−−

JN401094

HQ006966−−

JN401093

JN401092

JN401091

HQ006990

HQ006962

HQ007001

JN401090−

JN401089

JN401088

HQ006967

JN401087

JN401086−

HQ006987

GU828168

JN401730−

HQ006555

JN401743−

HQ006531

JN401746

JN401745

HQ006552−−

GU830093

JN401744

HQ006505

JN401742

JN401741

HQ006569−

HQ006515

JN401739

JN401738

JN401737

JN401736

JN401735

HQ006536

HQ006512

HQ006547

JN401734

HQ006509

JN401733

JN401732

HQ006516−

JN401731−−

GU830058

TS

TS

TG/TS

TG

TS

TS

TG/TS

TG/TS

TG

TG

TG

TG

TG

TS

TS

TS

TG/TS

TG

TS

TG

TG/TS

USA

GHANA

HONG KONG

HONG KONG

RUSSIAUAE

USA

HONG KONG

HONG KONG

MALAYSIA

HONG KONG

FINLANDFINLANDFINLANDGHANA

HONG KONG

HUNGARY

MALAYSIAINDONESIA

MALAYSIA

MALAYSIA

MALAYSIA

MALAYSIA

MALAYSIAHUNGARY

FINLAND

HONG KONG

MALAYSIA

HONG KONG

GHANA

GHANA

INDONESIAINDONESIAHONG KONG

HONG KONG

HONG KONG

FINLAND

Appendix
41

I,II
II
I,II
II
II
I,II
II
II
I,II
I,II
I,II
I,II
I,II
I,II
I,II
II
II
II
I,II
II
I,II
II
II
I,II,IV,V
I,II
II
II
II
II
I,II
I,II
I,II
I,II
I,II
I,II
II
II
I,II

Sypnoides fumosa
Daddala lucilla
Catephia alchymista
Heteranassa sp.
Zale bethunei
Thysania zenobia
Toxonprucha sp.
Pseudbarydia crespula
Pandesma robusta
Heteropalpia acrosticta
Sphingomorpha chlorea
Pericyma cruegeri
Sympis rufibasis
Erebus ephesperis
Erygia apicalis
Bulia deducta
Forsebia perlaeta
Melipotis punctifinis
Melipotis jucunda
Phoberia atomaris
Audea bipunctata
Audea humeralis
Hypotacha brandbergensis
Catocala sponsa
Ulotrichopus macula
Hypopyra capensis
Spirama retorta
Calyptis idonea
Ommatophora luminosa
Pantydia diemeni
Mocis latipes
Callistege mi
Euclidia glyphica
Ercheia cyllaria
Hulodes caranea
Ericeia subcinerea
Platyja umminea
Anisoneura salebrosa

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

RZ38

RZ261

RZ39

RZ126

RZ33

RZ82

MM05469

RZ20

RZ309

RZ407

RZ473

RZ359

RZ149

RZ241

MM04358

RZ275

RZ290

RZ60

RZ286

RZ58

RZ342

RZ284

RZ314

RZ29

RZ11

RZ48

RZ99

RZ291

RZ243

RZ321

RZ91

RZ307

RZ53

RZ270

RZ350

RZ127

RZ320

RZ313

HQ006213

JN401341

HQ006214

HQ006163

HQ006205

HQ006239

HQ006150

HQ006180

HQ006199

JN401340

JN401339

JN401338

HQ006172

HQ006185

GU828700

JN401337

JN401336

HQ006233

JN401335

HQ006230

JN401334

JN401333

JN401332

HQ006194

HQ006161

HQ006223

HQ006244

HQ006195

HQ006186

HQ006204

JN401331

JN401330

HQ006225

JN401329

JN401328

HQ006164

JN401327

HQ006201

HQ006917

JN401223

HQ006918−

HQ006909

HQ006942

HQ006857

HQ006884

HQ006903

JN401222

JN401221

JN401220

HQ006876

HQ006889

GU828495

JN401219

JN401218

HQ006936

JN401217

HQ006933

JN401216

JN401215

JN401214

HQ006898

HQ006866−

HQ006946

HQ006899

HQ006890

HQ006908

JN401213

JN401212

HQ006928

JN401211

JN401210

HQ006961

JN401209

HQ006905

HQ006309

JN401455

HQ006310

HQ006260

HQ006301

HQ006333

HQ006248

HQ006276

HQ006295

JN401454

JN401453

JN401452

HQ006269

HQ006281

GU829023

JN401451

JN401450

HQ006328

JN401449

HQ006325

JN401448

JN401447

JN401446

HQ006290

HQ006258

HQ006318

HQ006338

HQ006291

HQ006282

HQ006300

JN401445

JN401444

HQ006320

JN401443

JN401442

HQ006261

JN401441

HQ006297

HQ006400

JN401556

HQ006401−

HQ006394

HQ006426

HQ006343

HQ006371

HQ006388

JN401555−−

HQ006363−

GU829312

JN401554

JN401553

HQ006420−

HQ006417

JN401552

JN401551

JN401550

HQ006384

HQ006353

HQ006410

HQ006431

HQ006385

HQ006376

HQ006393

JN401549

JN401548

HQ006412

JN401547

JN401546

HQ006355

JN401545

HQ006390

HQ006821

JN401027

HQ006822

HQ006772

HQ006813

HQ006847

HQ006759

HQ006788

HQ006807

JN401026

JN401000

JN401025

HQ006781

HQ006793

GU829576

JN400999

JN401024

HQ006841

JN401023

HQ006838

JN401022

JN401021

JN401020

HQ006802

HQ006770

HQ006831

HQ006852

HQ006803

HQ006794

HQ006812

JN401019−

HQ006833

JN401018

JN401002

HQ006773

JN400998

HQ006809

HQ006476

JN401658

HQ006477

−

−

−

−

HQ006453

HQ006464

JN401657

JN401656

JN401655−

HQ006457

GU829816

JN401654

JN401653−−−

JN401652

JN401651

JN401650−

HQ006440

HQ006485

HQ006497−−−

JN401649

JN401648

HQ006486

JN401647

JN401646

HQ006441

JN401645

HQ006466

HQ006725

JN401960

HQ006726

HQ006679

HQ006718

HQ006750

HQ006667

HQ006695

HQ006712

JN401959

JN401958

JN401957

HQ006688

HQ006699

GU830688

JN401956

JN401955

HQ006744

JN401954

HQ006742

JN401953

JN401952

JN401951

HQ006707

HQ006677

HQ006735

HQ006755

HQ006708

HQ006700

HQ006717

JN401950

JN401949

HQ006737

JN401948

JN401947

HQ006680

JN401946

HQ006714−

JN401863

HQ006635

HQ006589

HQ006626

HQ006657

HQ006578

HQ006603

HQ006620

JN401862

JN401861−

HQ006596

HQ006608

GU830404−−

HQ006652

JN401859

HQ006650

JN401858

JN401857

JN401856

HQ006615

HQ006587

HQ006644

HQ006662

HQ006616−

HQ006625

JN401855

JN401854

HQ006645

JN401853

JN401860

HQ006590

JN401852

HQ006622

HQ007003−

HQ007004−

HQ006998

HQ007023

HQ006950

HQ006976

HQ006992

JN401108

JN401107

JN401106

HQ006968−

GU828189

JN401105

JN401104

HQ007019

JN401103

HQ007017

JN401102

JN401101

JN401100

HQ006988

HQ006959

HQ007012

HQ007027−

HQ006981

HQ006997

JN401099−

HQ007014−−

HQ006961

JN401098

HQ006994

HQ006548

JN401761

HQ006549−

HQ006543

HQ006570

HQ006500

HQ006525

HQ006538

JN401760

JN401759

JN401758

HQ006517

HQ006530

GU830086

JN401757

JN401756

HQ006565

JN401755

HQ006563

JN401754

JN401753

JN401752

HQ006535

HQ006510

HQ006557

HQ006575−−

HQ006542

JN401751

JN401750

HQ006558

JN401749

JN401748−

JN401747

HQ006539

TS

TS

TG

TG

TG

TS

TG/TS

TG

TG

TG

TG/TS

TS

TG/TS

TG

TS

TS

TG

TS

TG

TS

TG/TS

TG

TG/TS

JAPAN

HONG KONG

HONG KONG

HONG KONG

MALAYSIA

HONG KONG

FINLANDFINLANDCOSTA RICA

AUSTRALIA

MALAYSIAECUADORTAIWANGHANA

TAIWANFINLANDNAMIBIATANZANIA

CONGO

USA

USA

COSTA RICA

USA

USA

HONG KONG

TAIWANHONG KONG

HONG KONGTANZANIA

UAE

SPAIN

COSTA RICAUSA

COSTA RICA

USA

USA

GERMANY

JAPAN

42
Appendix

II
II
I,II
II
II
II
I,II
II
II
I,II
I,II,IV
II
I,II
I,II
IV

I,II,IV
IV

IV,V
IV,V
IV
IV
I,IV
IV
IV
IV
I,II,IV,V
IV
IV
IV

Ischyja manlia
Oxyodes scrobiculata
Serrodes campana
Avatha uloptera
Cocytia durvillii
Bastilla praetermissa
Achaea serva
Chalciope mygdon
Allotria elonymphaClytie devia
Ophiusa tirhaca
Thyas metaphaea
Ophiusa coronata
Artena dotata
Lepidodes gallopavo
Lepidodes limbulata 1
Lepidodes limbulata 2
Lepidodes limbulata 3
Diphthera festiva 2
Diphthera festiva 3
Diphthera festiva 1
Risoba obstructa 2
Risoba obstructa 1
Baileya levitans
Collomena siopera 1
Collomena siopera 2
Neostictopera nigropuncta
Beana terminigera
Baroa siamica
Selepa molybdea
Selepa discigera/celtis
Ptisciana seminivea
Triorbis aureovitta
Eligma narcissus
Elaeognatha argyritis
Iscadia productPterogonia cardinalis

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Erebinae

Diphtherinae

Diphtherinae

Diphtherinae

Diphtherinae

Diphtherinae

Diphtherinae

Diphtherinae

Risobinae

Risobinae

Risobinae

Collomeninae

Collomeninae

Collomeninae

Beaninae

Eligminae

Eligminae

Eligminae

Eligminae

Eligminae

Eligminae

Eligminae

Eligminae

Westermanniinae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Erebidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

IV

II,IV

IV

IV

IV

IV

IV

IV

II

Praxis porphyretica

Erebinae

Erebidae

RZ522

RZ361

RZ462

RZ97

RZ360

RZ516

RZ500

RZ32

RZ396

RZ638

RZ469

RZ634

RZ632

RZ461

RZ65

RZ381

RZ633

RZ631

RZ465

RZ594

X

X

X

HQ006243

X

X

X

HQ006203

X

X

X

X

X

X

X

JN401233

X

X

X

X

X

−

RZ630

X

RZ353

HQ006222

HQ006181

JN401350

HQ006187

HQ006188

JN401349

JN401348

HQ006179

JN401347

JN401346

JN401345

HQ006202

JN401344

JN401343

JN401342

RZ628

RZ46

RZ21

RZ190

RZ246

RZ247

RZ294

RZ391

RZ19

RZ306

RZ401

RZ317

RZ318

RZ113

RZ269

RZ308

X

X

X

HQ006945

X

X

X

HQ006907

X

−

X

−

−

X

X

JN401117

−

−

X

X

−

−

X

HQ006926

HQ006885

JN401232

HQ006891

HQ006892

JN401231

JN401230

HQ006883

JN401229

JN401228

JN401227

HQ006906

JN401226

JN401225

JN401224

X−

X

HQ006337

X

X

X

HQ006299

X

X

X

X

−

X

X

JN401351

X

−

X

−

X

X

X

HQ006317

HQ006277

JN401464

HQ006283

HQ006284

JN401463

JN401462

HQ006275

JN401461

JN401460

JN401459

HQ006298

JN401458

JN401457

JN401456

−

X

−

−

HQ006430

X

X

X

HQ006392

X

X

X

X

−

X

X

−

−

−

X

−

X

X

X

HQ006409

HQ006372

JN401564

HQ006377

HQ006378

JN401563

JN401562

HQ006370

JN401561

JN401560

JN401559

HQ006391

JN401558

JN401557

JN401028

X

X

X

HQ006851

X

X

X

HQ006811

X

X

X

X

X

X

X

JN400926

X

X

X

X

X

X

X

HQ006830

HQ006789

JN401034

HQ006795

HQ006796

JN401033

JN401001

HQ006787

JN401032

JN401031

JN401003

HQ006810

JN401030

JN401029

JN401659

X−

X

HQ006496

X

X−

HQ006468

X

X

X

X

−

X

X

JN401565

−

−

X

−

X

X

−

−

HQ006454

JN401667

HQ006458

HQ006459

JN401666

JN401665

HQ006452

JN401664

JN401663

JN401662

HQ006467

JN401661

JN401660

JN401961

X

X

X

HQ006754

X

X

X

HQ006716

X

X

X

X

X

X

X

JN401872

−

X

−

−

−

−

−

HQ006734

HQ006696

JN401969

HQ006701−

JN401968

JN401967

HQ006694

JN401966

JN401965

JN401964

HQ006715

JN401963

JN401962

JN401864

X

X

X

HQ006661

X

X

X

HQ006624

X

X

X

X

X

X

X

JN401771

−

−

X

−

X

X

X

HQ006643

HQ006604

JN401871

HQ006609

HQ006610

JN401870

JN401865

HQ006602

JN401868

JN401869

JN401867

HQ006623

JN401866

JN401822

JN401109

X−−

HQ007026

X−

X

HQ006996

X

X

X

−

−

X

−

JN401035

−

−

X

−

X

X

X

HQ007011

HQ006977

JN401116

HQ006982

HQ006983

JN401115

JN401881

HQ006975

JN401113

JN401112−

HQ006995

JN401111

JN401110

JN401762−−−

HQ006574

X

X

X

HQ006541

X

X

X

−

−

X

X

JN401668

−

−

X

−

X

X

−

HQ006556

HQ006526

JN401770−−

JN401769

JN401768

HQ006524

JN401767

JN401766

JN401765

HQ006540

JN401764

JN401763

TS

TG/TS

TS

TG

TS

TG

TG

TG

TG

TG/TS

TG/TS

TG/TS

TS

TS

TS

TG

TG/TS

TS

TS

TG/TS

TS

TS

AUSTRALIA

MALAYSIACOSTA RICA

COSTA RICA

HONG KONG

MALAYSIA

MALAYSIA

HONG KONG

HONG KONG

MALAYSIATHAILANDCOSTA RICA

COSTA RICA

COSTA RICAUSA

THAILANDMALAYSIACOSTA RICA

COSTA RICA

COSTA RICA

COSTA RICA

COSTA RICA

COSTA RICA

COSTA RICA

HONG KONG

MALAYSIA

GHANA

UAE

UAE

USA

MALAYSIA

MALAYSIA

MALAYSIANEW GUINEA

MALAYSIATAIWANHONG KONG

MALAYSIA

Appendix
43

IV
IV
I,II,IV
IV

IV

Westermannia superba
Iragaodes nobilis
Negeta contrariata 2
Negeta abbreviata
Negeta contrariata 1
Negeta contrariata 3

Westermanniinae

Westermanniinae

Westermanniinae

Westermanniinae

Westermanniinae

Westermanniinae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae
IV
IV
IV
IV
IV
IV
IV
I,II,IV
IV
IV
IV
IV
IV
IV
IV
I,IV
IV
I,II,IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
I,IV
IV

Melanographia flexilineata
Membranola lampang
Alcanola tympanistes
Alcanola obscurata
Meganola minuscula
Nola basinigra
unknown noline
Nola aerugula
Nola minna
Nola cretacea
Nola analis
Nola lucidalis
Nola fasciata
Manoba brunellus
Maceda mansueta
Paracrama angulata
Paracrama dulcissima
Earias clorana
Earias roseifera
Earias insulana
Maurilia iconica
Tortriciforma razowskii
Didigua viridifascia
Xenochroa annae
Xenochroa xanthia
Diehlea tumida
Chloriola gratissima
Gelastocera viridimacula
Beara tortriciformis
Ariolica argentea
Gariga mirabilis

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Nolinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

NolidaeIV

IV

Miaromima cornucopia

Westermanniinae

Nolidae

RZ114

RZ63

RZ515

RZ521

RZ531

RZ479

RZ503

RZ386

RZ525

RZ586

RZ484

RZ492

RZ480

MM06650

RZ648

RZ43

RZ641

RZ495

RZ534

RZ533

RZ494

RZ493

RZ488

MM01776

RZ526

RZ627

RZ485

RZ625

RZ624

RZ626

RZ498

RZ504

RZ487

RZ414

RZ26

RZ483

RZ645

RZ530

X

HQ006234

X

X

X

X

X

X

X

X

X

X

X

GU828747

X

HQ006219

X

X

X

X

X

X

X

GU828670

X

X

X

X

X

X

X

X

X

X

HQ006191

X

X

X

X

HQ006937

X

X

X

X

X

X

X

X

X

X

X

GU929721−

HQ006923

−

X

−

X

X

X

X

GU828468

X

X

X

−

X

X

X

X

X

−

HQ006895

X

−

X

X

HQ006329

X

X

X

X

X

X

X

X

−

X

X

GU829061

X

HQ006315

−

X

X

X

−

X

X

GU829001

X

X

X

X

X

X

X

X

X

X

HQ006287

X

X

X

−

HQ006421

X

X

X

X

X

X

X

X

X

−

X

GU829346

X

HQ006406

−

X

X

X

X

−

X

GU829284

X

X

X

X

X

X

X

X

−

X

HQ006381

X

X

X

X

HQ006842

X

X

X

X

X

X

X

X

X

X

X

GU829610

X

HQ006827

X

X

X

X

X

X

X

GU829555

X

X

X

X

X

X

X

X

X

X

HQ006799

X

X

X

X

−

X

X

X

−

X

X

X

X

X

X

X

GU829845

X

HQ006482

−

X

X

−

X

X

X

−

X

X

X

X

X

X

X

X

X

X

HQ006460

X

X

X

X

HQ006745

X

X

X

X

X

X

X

X

X

−

X

GU830718

X

HQ006731

X

−

X

X

−

X

X

GU830665

X

X

X

X

X

X

X

X

X

X

HQ006704

X

X

X

X

HQ006653

X

X

X

X

X

X

X

X

X

X

X

GU830448

X

HQ006640

X

X

X

X

−

−

X

GU830376

−

X

−

X

X

X

X

X

X

X

HQ006612

−

X

X

−

−

HQ007020

−

−

X

X

X

X

−

−

X

X

X

GU828231−

HQ007008

−

X

−

−

X

X

−

GU828167

X

X

−

X

X

−

X

X

−

X

HQ006986−

X−

HQ006566

X

X

X

−

X

X

X

−

X

X

X

GU830131−

HQ006553

−

X

X

−

−

−

X

GU830057

X

X

X

−

X

X

X

X

X

X

HQ006533

−

−

X

TG

TG

TS

TS

TG

TG/TS

TS

TS

TG

TG

TG

TG

TG

TG

TG

TS

TS

TS

TG/TS

MALAYSIA

HONG KONG

JAPAN

MALAYSIA

MALAYSIA

MALAYSIA

MALAYSIA

MALAYSIA

MALAYSIA

MALAYSIA

MALAYSIAINDONESIAHONG KONG

TAIWANFINLANDTHAILAND

HONG KONG

THAILAND

HONG KONG

MALAYSIA

MALAYSIA

HONG KONG

HONG KONG

USA

FINLAND

MALAYSIATHAILANDUSA

THAILANDTHAILANDTHAILAND

HONG KONG

MALAYSIATAIWANMALAYSIA

HONG KONGJAPAN

THAILAND

44
AppendixI,IV,V
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
I,IV
IV
IV
IV
I,IV
IV
IV
IV
IV
I,IV
IV
IV
IV
IV
IV

IV
I,II,V
II,V
I,II,V
V
II,V

Pseudoips prasinana 1
Clethrophora distincta
Hylophilodes orientalis
Chloroplaga sp.
Chloroplaga nygmia
Sinna extrema
Sinna floralis
Siglophora ferreilutea
Chandica quadripennis
Cossedia hyriodes
Labanda ceylusalis
Aquis orbicularis
Tathothripa continua
Blenina donans
Blenina quinaria
Dilophothripa chrysorrhaea
Egchiretes nomimus
Nycteola degenerana
Nycteola sinuosa
Nycteola indica
Garella nilotica
Garella ruficirra
Giaura robusta
Giaura nigrilineata
Giaura multipunctata
Giaura niveidisca
Pardasena verna
Etanna clopaea
Etanna breviuscula 2
Etanna breviuscula 1
Etanna breviuscula 3
Dyops chromatophila
Pseudoarcte melanis
Arcte modesta
Pararcte schneideriana
Belciana kala

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Chloephorinae

Dyopsinae

Dyopsinae

Dyopsinae

Dyopsinae

Dyopsinae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

Nolidae

NolidaeNoctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

IV

IV

IV

Pseudoips prasinana 2

Chloephorinae

NolidaeIV

Tympanistes fusimargo

Chloephorinae

Nolidae

RZ416

RZ548

RZ54

RZ147

RZ10

RZ518

RZ497

RZ491

RZ486

RZ532

RZ527

RZ496

RZ519

RZ31

RZ502

RZ477

RZ539

RZ501

MM00135

RZ481

RZ528

RZ499

RZ64

RZ640

RZ639

RZ555

RZ524

RZ579

RZ489

RZ529

RZ490

RZ646

RZ520

RZ643

RZ478

MM00107

RZ128

RZ566

JN401238

X

HQ006226

JN401235

HQ006158

X

X

X

X

X

X

X

X

HQ006200

X

X

X

X

GU828612

X

X

X

HQ006235

−

X

X

X

X

X

X

X

X

X

X

X

GU828600

X

X

−

X

HQ006929

JN401119

−

X

X

X

X

X

X

X

−

HQ006904

X

X

X

X

GU828410

X

X

X

HQ006938

−

−

X

X

X

X

X

X

−

X

−

X

GU828399−

X

JN401356

X

HQ006321

JN401353

HQ006255

X

X

X

X

X

X

X

X

HQ006296

X

X

X

X

GU828946

X

X

X

HQ006330

X

X

X

X

X

X

X

X

X

X

X

X

GU828934

X

X

JN401468

X

HQ006413

JN401465

HQ006350

X

X

X

X

X

X

X

X

HQ006389

X

X

X

X

GU829238

X

X

X

HQ006422

X

X

X

X

X

X

X

X

X

X

X

X

GU829229

X

X

JN400930

X

HQ006834

JN400928

HQ006767

X

X

X

X

X

X

X

X

HQ006808

X

X

X

X

GU829506

X

X

X

HQ006843

X

X

X

X

X

X

X

X

X

X

X

X

GU829496

X

X

JN401570

X−

JN401567

HQ006438

X

X

X

X

−

X

X

X

HQ006465

−

X

X

X

GU829760−

X

X

HQ006490

X

X

X

X

X

X

X

X

X

X

−

X

GU829754

X

X

JN401879

X

HQ006738

JN401874

HQ006674

X

X

X

X

X

X

X

X

HQ006713

X

X

X

X

GU830621

X

X

X

HQ006746

X

X

X

X

X

X

X

X

X

X

X

X

GU830611

X

X

JN401776

X

HQ006646

JN401773

HQ006585

X

X

X

X

X

X

X

X

HQ006621

X

X

X

X

GU830323−

X

X

HQ006654

X

X

X

X

X

X

X

X

X

X

X

X

GU830312−X

JN401040−

HQ007015

JN401037

HQ006957

−

−

X

−

X

−

X

−

HQ006993

−

X

−

−

GU828113−−X

HQ007021

−

−

−

−

−

X

−

X

−

X

X

X

GU828101−X

JN401672−

HQ006559

JN401669

HQ006507

X

X

X

X

−

X

X

X

−

X

X

X

−

GU830000−−

X

HQ006567

−

−

X

−

X

−

X

X

X

X

−

X

GU829989

−

−

TS

TG

TG

TG

TG

TG

TS

TS

TG

TG

TS

TS

TS

TS

TS

TG

TG/TS

MALAYSIACOSTA RICA

MALAYSIAGHANACOSTA RICA

MALAYSIA

HONG KONG

HONG KONG

AUSTRALIA

MALAYSIA

MALAYSIA

HONG KONG

MALAYSIA

HONG KONG

HONG KONG

USA

MALAYSIA

HONG KONG

FINLANDCOSTA RICA

MALAYSIA

HONG KONG

SUMATRATHAILANDTHAILANDMALAYSIA

MALAYSIA

MALAYSIAJAPAN

MALAYSIAJAPANTHAILANDMALAYSIATHAILANDJAPAN

FINLANDGERMANYMALAYSIA

Appendix
45

II,V
V
II,IV,V
II,IV,V
II,IV,V
IV,V
IV,V
IV,V
I,II,IV,V
I,V
V
I,V

Encruphion leena
Vespola caeruleifera
Parangitia temperata
Parangitia mosaica
Diopa corone
Concana mundissima
Concana lecta
Concana permixta
Amphipyra perflua
Brachionycha nubeculosa
Flammona quadrifasciata
Panemeria tenebrataBagisarinae

Bagisarinae

Bagisarinae

Bagisarinae

Bagisarinae

Bagisarinae

Bagisarinae

Bagisarinae

Amphipyrinae

Amphipyrinae

Metoponiinae

Metoponiinae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

VII,V

Dyrzela plagiata

Bagisarinae

Noctuidae

V

II,V

Ramadasa pavo

Bagisarinae

Noctuidae

Comachara cadburyi

I,II,V

Xanthodes albago

Bagisarinae

Noctuidae

Cerma cerintha

II,VAmyna octo

Bagisarinae

Noctuidae

Acronictinae

I,IV,V

Acontia lucida

Acontiinae

Noctuidae

Acronictinae

I,VEmmelia trabealis

Acontiinae

Noctuidae

Noctuidae

I,II,V

Panthea coenobita

Pantheinae

Noctuidae

Noctuidae

II,V

Antitrisuloides catocalina

Pantheinae

Noctuidae

I,V

II,V

Thiacidas sp.

Pantheinae

Noctuidae

Acronicta rumicis

II,V

Eucocytia meeki

Pantheinae

Noctuidae

Acronictinae

I,II,V

Raphia abrupt

Raphiinae

Noctuidae

Noctuidae

II,V

Diloba caeruleocephala

Dilobinae

Noctuidae

V

I,IV,V

Cucullia umbratica

Cuculliinae

Noctuidae

I,V

I,V

Deltote uncula

Eustroniinae

Noctuidae

Craniophora ligustri

I,V

Abrostola tripartita

Plusiinae

Noctuidae

Acronicta americana

I,II,IV,V

Autographa gamma

Plusiinae

Noctuidae

Acronictinae

II,V

Sosxetra grata

Dyopsinae

Noctuidae

Acronictinae

V

Ceroctena amynta

Dyopsinae

Noctuidae

Noctuidae

V

Cyclodes omma

Dyopsinae

Noctuidae

Noctuidae

II,V

Belciana biformis

Dyopsinae

Noctuidae

RZ618

RZ617

MM01529

RZ597

MM06745MM00005

RZ596

MM01542MM01162

RZ482

RZ474

RZ621

RZ472

RZ464

RZ463

RZ595

RZ351

RZ395

RZ382

MM09894

RZ50

MM00152

MM09893MM04583

RZ388

RZ459

RZ87

CWM-94-0372

MM09267

MM04543

MM04601

MM05132

MM00328

RZ281

RZ552

RZ636

RZ384

X

X

GU828666

X

HQ006148

HQ006157

X

GU828667

GU828660

X

X

X

JN401239

JN401245

JN401244X

JN401243

JN401240

JN401241

GU828844

JN401242

GU828617

HQ006147

GU828702

JN401248

JN401249

JN401247

GU828548

JN401246

GU828701

GU828703

HQ006152GU828636

JN401236

X

X

JN401237

X

X

GU828464

X

HQ006855

HQ006863

X

GU828465

GU828458

X−

X

JN401122−

JN401127X

JN401126

JN401123

JN401124

GU929808

JN401125

GU828415

HQ006854

GU828497

JN401130

JN401131

JN401129

GU828350

JN401128

GU828496

GU828498

HQ006859

GU828434

JN401120XX

JN401121

X

X

GU828997

X

HQ006246

HQ006254

X

GU828998

GU828991

X−

X

JN401357

JN401363

JN401362

X

JN401361

JN401358

JN401359

GU829145

JN401360

GU828952

HQ006245

GU829025

JN401365

JN401366−

GU828893

JN401364

GU829024

GU829026

HQ006250

GU828970

JN401354

X

X

JN401355

X

X

GU829280

X

HQ006341

HQ006349

X

GU829281

GU829275

X

X

X

−

JN401474

JN401473

X

JN401472

JN401469

JN401470

GU829412

JN401471

GU829243

HQ006340

GU829314

JN401476

JN401477−

GU829193

JN401475

GU829313

GU829315

HQ006345

GU829256

JN401466

X

X

JN401467

X

X

GU829551

X

HQ006757

HQ006766

X

GU829552

GU829546

X

X

X

JN400931

JN400935

JN400934

X

JN400933

JN400932

JN400978GU829693

JN400984

GU829512

HQ006756

GU829578

JN400936

JN400937−

GU829455

JN400982

GU829577

GU829579

HQ006761

GU829528

JN400988

X

X

JN400929

X

X

GU829792

X

HQ006432

HQ006437

X

GU829793

GU829787

X−

X

JN401571

JN401577

JN401576−

JN401575

JN401572

JN401573−

JN401574

GU829763−−

JN401578

JN401579−

GU829728−

GU829817

GU829818−−

JN401568

X

X

JN401569

X

X

GU830662

X

HQ006665

HQ006673

X

GU830663

GU830657

X

X

X

JN401880

JN401886

JN401885

X

JN401884

JN401881

JN401882

GU830792

JN401883

GU830627

HQ006664

GU830690

JN401889

JN401888

JN401888

GU830579

JN401887

GU830689

GU830691

HQ006669

GU830640

JN401875

X

X

JN401877

X−

GU830372

X

HQ006577

HQ006584

X

GU830373

GU830366

X

X−

JN401777

JN401783

JN401782

X

JN401781

JN401778

JN401779

GU830535

JN401780

GU830327

HQ006576

GU830406

JN401786

JN401787

JN401785

GU830270

JN401784

GU830405

GU830407

HQ006580

GU830344

JN401774

X

X

JN401775−−

GU828163−

HQ006948

HQ006956

X

GU828164

GU828157−−−

JN401041

JN401046

JN401045

X

JN401044

JN401042−

GU828308

JN401043

GU828118

HQ006947

GU828191

JN401048−

JN401047

GU828059−

GU828190

GU828192

HQ006952

GU828135

JN401038

X−

JN401039

X

X

GU830053

X

HQ006498

HQ006506

X

GU830054

GU830047

X

−

X

−

JN401677−−

JN401676

JN401673

JN401674

GU830224

JN401675

GU830006−

GU830088

JN401679

JN401680−

GU829939

JN401678

GU830087

GU830089

HQ006502

GU830023

JN401670−

X

JN401671

TS

TG

TG

TS

TS

TS

TG

TS

TS

TS

TS

TG

TG/TS

TG

TG

TG/TS

TG

TG/TS

TG/TS

TG

TG/TS

TS

TS

TS

TS

USA

USA

FINLANDUSAFINLANDFINLAND

MALAYSIAFINLANDFINLANDGUATEMALACOSTA RICAUSA

FRENCHGUIANACOSTA RICA

COSTA RICA

COSTA RICA

COSTA RICA

MALAYSIA

MALAYSIA

SARDINIA

MALAYSIAGREECE

SARDINIAFINLAND

MALAYSIA

INDIA

INDONESIAUSAFINLANDFINLANDFINLANDFINLANDFINLANDCOSTA RICA

ECUADORTHAILANDMALAYSIA

46
AppendixV
V
V
I,II,V
II,V
V
I,II
II,V
I,V
V
I,V
V
V
I,II,V
V
I,V
I,V
I,II,V
I,IV,V
I,V
I,II,IV,V

Harrisimemna trisignata
Polygrammate hebraeicum
Agriopodes fallax
Periscepta polysticta
Aedia leucomelas
Condica illecta
Condica vecors
Hemicephalis alesa
Pyrrhia umbra
Chytonix diehli
Cryphia raptricula
Stenoloba futii
Ecpatia sp.
Ecpatia longinquua
Tiracola aureata
Actinotia polyodon
Hoplodrina octogenaria
Diaphone sp.
Apamea crenata
Ufeus faunas
Noctua fimbriata

Acronictinae

Acronictinae

Acronictinae

Agaristinae

Aediinae

Condicinae

Condicinae

Condicinae

Heliothinae

unassigned

Bryophilinae

Bryophilinae

unassigned

unassigned

Noctuinae

Noctuinae

Noctuinae

Noctuinae

Noctuinae

Noctuinae

Noctuinae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

Noctuidae

MM04752

RR-98-0914

MM01170MF-05-0053

MM01651

MM05153

RZ14

RZ25

RZ545

RZ523

MM04919

RZ517

MM05114

RZ341

CWM-95-0471

RZ511

RZ277

MM07669

RZ616

RZ620

RZ619

GU828705

GU828860

GU828661

GU828571

HQ006153

GU828714

X

HQ006190

X

X

GU828708

X

GU828712

JN401251

GU828550

X

JN401250

GU828820

X

X

X

GU828500

GU929822

GU828459

GU828372

HQ006860

GU828509

X

HQ006894

X

X

GU828503

X

GU828507

JN401133

GU828352

X

JN401132

GU929788

X

X

X

GU829028

GU829163

GU828992

GU828913

HQ006251−

X

HQ006286

X

X

GU829031

X

GU829034

JN401368

GU828895

X

JN401367

GU829125

X

X

X

GU829317

GU829425

GU829276

GU829206

HQ006346

GU829326

X

HQ006380

X

X

GU829320

X

GU829324

JN401479

GU829194

X

JN401478

GU829400

X

X

X

GU829581

GU829709

GU829547

GU829475

HQ006762

GU829590−

HQ006798

X

X

GU829584

X

GU829588

JN400938

GU829457

X

JN400976

GU829674

X

X

X

GU829820

GU829911

GU829788

GU829738

HQ006434

GU829827

X

−

X

X

GU829822

X

GU829825

JN401581−

X

JN401580

GU829892

X

X

X

GU830693

GU830807

GU830658

GU830591

HQ006670

GU830702

X

HQ006703

X

X

GU830696

X

GU830700

JN401892

GU830581

X

JN401891

GU830773

X

X

X

GU830409

GU830552

GU830367

GU830285

HQ006581

GU830418

X

HQ006611

X

X

GU830412

X

GU830416

JN401789−−

JN401788

GU830519

X

X

X

−

GU828194

GU828320

GU828158

GU828076

HQ006953

GU828202X

HQ006985−

X

GU828196−

GU828200

JN401049

GU828061−−

GU828289−

X

X

GU830091

GU830238

GU830048

GU829960

HQ006503

GU830100−

HQ006532

X−

GU830094

X

GU830098

JN401682

GU829941

X

JN401681

GU830201

X

X

TG

TG

TG

TG

TG

TG

TS

TG/TS

TS

TS

USA

FINLANDUSAFINLANDTANZANIAFINLANDFINLAND

INDONESIAHONG KONG

MALAYSIA

MALAYSIAFINLAND

MALAYSIAFINLANDCOSTA RICAUSA

MALAYSIAJAPAN

AUSTRALIA

USA

USA

Appendix47