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TURUN YLIOPISTON JULKAISUJAANNALES UNIVERSITATIS TURKUENSIS
SARJA - SER. AII OSA - TOM. 268
BIOLOGICA - GEOGRAPHICA - GEOLOGICA
TURUN YLIOPISTOUNIVERSITY OF TURKU
Turku 2012
Molecular SySteMaticS
of Noctuoidea (iNSecta, lepidoptera)
reza zahiri
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
Finland
Co-advised by: Ph.D. J. Donald Lafontaine Canadian National Collection of Insects, Arachnids and Nematodes Canada Ph.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 Zahiri
This thesis is based on the following original research contributions, which are referred to in 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 a
stable 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 their
fruit-piercing relatives (Lepidoptera: Erebidae: Calpinae). Submitted
manuscript
IV 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 of
Noctuidae (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 a quarter 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 for the superfamily, then build up robust phylogenetic frameworks for those circumscribed monophyletic entities (i.e., families), as well as clarifying the internal classification of monophyletic 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 well-resolved molecular phylogenetic hypothesis for higher taxa of Noctuoidea consisting of six strongly supported families: Oenosandridae, Notodontidae, Euteliidae, Erebidae, Nolidae, and Noctuidae. The studies in my thesis highlight the importance of molecular data in systematic and phylogenetic studies, in particular DNA sequences of nuclear genes, and an extensive sampling strategy 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 Why Noctuoidea? .................................................................................................... 7 1.2 Status of Noctuoidea ................................................................................................ 8 1.3 Initiation of molecular phylogenetic approaches ................................................... 10 1.4 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 Phylogenetic hypothesis for Noctuoidea ............................................................... 16 3.2 Pattern of relationships among major lineages of Noctuoidea .............................. 17 3.3 Phylogenetic hypothesis for quadrifid Noctuoidea ................................................ 19 3.4 Character optimizations ......................................................................................... 21 3.5 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 explains various large and independent classes of facts, it rises to the rank of a well-grounded theory”
Charles Darwin 1. INTRODUCTION
1.1 Why Noctuoidea?
I quote a magnificent passage from Charles Darwin who said ‘‘all the organic beings which have ever lived on this earth have descended from some one primordial form.’’ 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 level to 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, and insects 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 and butterflies) is one of four super-radiations of insects (along with beetles, flies and wasps) that account for the majority of animal life on Earth. Noctuoidea are the largest superfamily within Lepidoptera—belonging to a large ditrysian clade that also includes e.g., geometroids and bombycoids (Regier et al., 2009, Mutanen et al., 2010)—with approximately 45,000 described (Nieukerken et al., 2011) and
many unknown as well as unnamed species, particularly from tropical regions.
To understand Noctuoidea evolution, their systems of evolutionary relationships—phylogenies based on extensive evidence from living lineages of noctuoids—must be recognized. Fortunately, the monophyly of Noctuoidea is firmly established. It is based on the presence of a single apomorphic character, the metathoracic tympanal organ (Miller, 1991). This organ is a highly specialized hearing apparatus that detects the echolocation signals of bats (Kitching & Rawlins, 1998); however, there is increasing evidence that the tympanum may also be involved in reception of mating signals (Kitching & Rawlins, 1998).
Noctuoidea, like most lepidopterans, are plant feeding as caterpillars and nectar feeding as adults, and they are a prominent element of terrestrial ecosystems, functioning as herbivores, pollinators and prey, as well as being one of the most damaging 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
Introduction
8
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, the adults of some genera damage fruit crops by piercing the skins to suck juices (Baenziger, 1982).
The study of relationships among major lineages of organisms—phylogenetic analysis—has been closely associated with the classification and naming of the diversity of life on Earth (i.e., taxonomy), which both are a branch of the science of systematics. The classification of organisms has been one of the major ongoing accomplishments of human society (Wilson, 2000), but is still far from completion, both in terms of the inventory of species and of the classification of those species in a hierarchical system that has a phylogenetic basis. 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, in particular DNA sequences, is becoming increasingly important for testing and improving classifications, especially for highly diverse groups of organisms such as insects.
The main theme of my thesis, as well as my ongoing research, is to improve our understanding of the reasons and causes behind the flourishing diversification of Noctuoidea on Earth. To address tens of such questions—diversity of their adaptations, biomass, species-richness, ecological and economical impacts, etc.—it is necessary, as a first step, to place Noctuoidea and its major lineages in a phylogenetic context, by reconstructing a
strong phylogenetic hypothesis. The resolution of a stable, extrapolative higher-level classificatory structure for the major lineages of Noctuoidea, and understanding their phylogenetic relationships, is also of importance 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 the fundamental distinction between the different systems is based on the use of unsatisfactory (occasionally plesiomorphic) characters in phylogenetic reconstruction. Various authors have recognized between five and thirteen families, and strikingly, no two publications have agreed on the same divisions of the superfamily into families (Kitching & Rawlins, 1998, Lafontaine & Fibiger, 2006). For instance, in Figure 1, I have summarized most recent Noctuoidea classifications and have compared them with the most recent one, which is presented in this thesis (I). Kitching (1984) published a historical review of noctuid subfamily relationships and showed that the higher classifications of Noctuidae used up to that time had been based upon superficial resemblance and vaguely defined characters, rather than on rigorous application of cladistic principles. Subsequently, Speidel and co-workers attempted to progress beyond the age of traditional morphological noctuid taxonomy by initiating investigations based mainly on the male genitalia and the tympanal region, which they considered particularly 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 of the systematics of noctuoids, and made it clear that increased character and taxon sampling were necessary to resolve the relationships 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), each
classification having its own limitations and strengths (Roe et al., 2010). Fibiger & Lafontaine (2005) proposed a rather new classification with ten families: Oenosandridae, Doidae, Notodontidae, Strepsimanidae, Nolidae, Lymantriidae, Arctiidae, Erebidae, Micronoctuidae and Noctuidae (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
Introduction
10
and Erebidae sensu Fibiger & Lafontaine (2005) were downgraded to subfamily status within an expanded family concept of Noctuidae based on the quadrifid venation of the forewing and the presence of a tympanal sclerite in the tympanal membrane. In their view, the superfamily should consist of five families: Oenosandridae, Doidae, Notodontidae, Micronoctuidae and Noctuidae (Figure 1).
To resolve the dominant complexity of relationships in higher systematics of Noctuoidea, it was crucial to understand their phylogenetic relationships by building a robust skeleton phylogenetic hypothesis, i.e., a phylogeny that was based on certain specialized features, and that had a common ancestor and unique evolutionary history. To achieve a more robust phylogenetic hypothesis, one strategy is to increase the number of characters to obtain a dataset with a strong phylogenetic signal. In molecular systematics, 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 and heterogeneous group such as the Noctuoidea (Speidel & Naumann, 1995) with a vast number of species presents obstacles to the use of morphology at this taxonomic level. The adults and larvae of species in Noctuoidea exhibit a bewildering 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 states
correctly and can be subject to extensive homoplasy (character convergence and reversal). As a result, morphological analyses have often failed to determine relationships among most groups with confidence.
1.3 Initiation of molecular phylogenetic approaches
As noted above, until recently, the higher systematics of Noctuoidea had been based primarily on morphological characters with a predominantly phenetic approach, until the introduction and application of cladistic philosophy by Kitching (1984, 1987) and Miller (1991). More recently, molecular data, in particular DNA sequences of mitochondrial and nuclear genes (mtDNA and nDNA, respectively), have opened new and fruitful avenues for the study of phylogenetic relationships. Molecular studies 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, and even entire mitochondrial genomes (15,000–20,000 bp in insects) fail to provide robust inferences at deep levels (Cameron et al., 2004). However, they are useful 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 mid-Tertiary and later age, although they have also been applied, albeit infrequently, at deeper levels (Wiegmann et al., 2000). In contrast, protein-coding nuclear genes generally have slower mutation rates compared with mtDNA. They are most
Introduction
11
frequently used to study older evolutionary divergences and are particularly good at resolving 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). Over the past few years, a series of papers have been published with a shared objective: to contribute to the development of a more satisfactory classification of Noctuoidea at levels above that of the genus and particularly the ‘quadrifid’ part—those noctuoids with forewing vein M2 arises closer to the origin of M3 than M1, in the lower part of the discal cell—of the superfamily. It began with the exploration of 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 old concept of Noctuidae (Mitchell et al., 1997). Mitchell et al. (2006) found a strongly supported clade of quadrifine noctuid moths that also included the families Lymantriidae and Arctiidae. They termed this 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 enigmatic family Doidae did not group with the other noctuoids, but instead appeared to be related to Drepanoidea (Regier et al., 2009, Mutanen et al., 2010). Otherwise both studies found Noctuoidea to be monophyletic, with Oenosandridae being sister to the rest and Notodontidae the next lineage branching off.
However, all of these studies had very poor sampling of the higher taxa putatively belonging to the L.A.Q. clade, and critically they did not sample type genera of many higher taxa. Given that the monophyly of many named groups remains questionable, it is crucial to sample the type genera of each family, subfamily and tribe to assess the taxonomic limits of a given category. Furthermore, previous molecular studies have used only a small number of molecular markers, usually one to three gene regions (Wahlberg & Wheat, 2008).
1.4 Outline of the thesis
In this thesis, I employed the methods of molecular phylogenetics using eight markers 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 a comprehensive overview of the higher-level 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 are interpreted as families (I). The second chapter aims to understand the higher-level phylogeny and elucidate the evolutionary history of the moth family Erebidae, the most controversial group among the newly circumscribed families (II). Erebidae is a massive clade and includes a diverse groups exhibiting a broad range of feeding behaviors, including those that can be considered ‘piercers’ of fruits or other hosts (skin-piercers: hematophagous) and ‘tear feeders’ (lachryphagous) (III). Within butterflies and moths, adult hematophagy is limited to species within the vampire moth genus Calyptra
Introduction
12
Ochsenheimer, which are placed within the subfamily Calpinae, Erebidae. Paper III focused on the subfamily Calpinae using both morphological and molecular data to reconstruct ancestral feeding behaviors within Calpinae as well as whether fruit-piercing behavior and associated modifications of the tongue have evolved independently in different groups of Erebidae (III). The fourth chapter seeks to elucidate the deep divergences and evolutionary 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 of previously uncertain affinity—was recovered. Diphtherinae is considered to be the plesiomorphic sister lineage to the rest of Nolidae (IV), thus a new phylogenetic hypothesis for Nolidae is presented (IV). The fifth chapter elucidates the evolutionary 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 their main 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 to include representatives of all known major lineages of quadrifine Noctuoidea from every biogeographic region. This strategy is being used to establish priorities for ongoing studies, to test further the robustness of the major clades of Noctuoidea, both in relation to each other and 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 most recognized Noctuoidea lineages for papers I–V, respectively. Furthermore, there were numerous unplaced taxa of uncertain status that were included in each paper. Indeed, the allocation of the unplaced taxa into any higher taxa of Noctuoidea, however tentatively, could not be determined in previous classifications. I was unable to sample/amplify a few rare taxa with restricted distributions and/or low species richness (e.g., Strepsimaninae, Afridinae, Camptolominae). Where possible, a representative of the type genus of each lineage is included, but this was not possible for a few tribes/subtribes, in which 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 most
closely related taxa of other families of Lepidoptera (papers I, II, IV, V) or subfamilies of Erebidae (III). I rooted the cladograms, in different papers, with different taxa, which represents what I consider to be the putative sister family to the 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, and elongation factor-1α (EF-1α), ribosomal protein S5 (RpS5), carbamoylphosphate synthase domain protein (CAD), cytosolic malate dehydrogenase (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, protein-coding exons and have previously been found 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) and sequencing protocols follow Wahlberg & Wheat (2008). Sequencing was performed mainly with an ABI 3730XL capillary sequencer (Macrogen, Seoul, Korea), and a smaller part with an ABI PRISMR 3130XL capillary sequencer (Turku,
Material and Methods
14
Finland). The resulting chromatograms were checked and DNA sequences aligned by 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 neighbor-joining and Maximum Likelihood trees separately for each gene region and checked them carefully for identical sequences and other doubtful patterns. In addition, to minimize the risk of misidentification, all the specimens were cross-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 were available for many of the species used in this 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 non model-based (i.e., parsimony) methods. Initially, the data matrices were analysed in 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 was partitioned into mtDNA and nDNA and finally the data was partitioned by gene regions (8 partitions). The effects of varying taxon and gene combinations were compared 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 the data by genes (8 partitions) in the ML analyses, and by nDNA and mtDNA (two partitions) 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, with default parameters applied, until the minimal tree was found 1000 times. All characters were treated as unordered and equally weighted, and robustness of the hypothesis was assessed 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 of sequence evolution for each gene partition based on the Akaike Information Criterion using FindModel (http://www.hiv.lanl.gov/ content/sequence/findmodel/findmodel.html). 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 carried out 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
Material and Methods
15
mixing of chains and problems with convergence of likelihoods. I modeled the evolution of sequences according to the GTR + Γ model independently for the two partitions using the “unlink” command in MrBayes. The Bayesian analyses were separately run two times for five, 23 and 20 million generations for papers III−V, respectively, with every 1000th generation sampled. Clade robustness was estimated by posterior probabilities (PP) in MrBayes. Convergence was determined when the standard deviation of split frequencies went below 0.05 and the PSRF (Potential Scale Reduction Factor) approached 1, and both runs had properly converged to a stationary distribution after the burn-in
stage (which was 1,000 sampled generations).
To understand character evolution in higher 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 and character transformations were optimized onto the topology resulting from the Bayesian analysis (IV).
All laboratory procedures and phylogenetic data analyses are detailed in the original papers.
Results and Discussion
16
3. RESULTS AND DISCUSSION
3.1 Phylogenetic hypothesis for Noctuoidea
In the initial study, I aimed, as a first step, to reconstruct a robust phylogenetic hypothesis for higher taxa of Noctuoidea. The results strongly supported the monophyly of Noctuoidea. The major groups within the Noctuoidea clade all shared a particular morphological synapomorphy—a metathoracic tympanal organ. I found six strongly supported major lineages within Noctuoidea that deserved family status. These are Oenosandridae, Notodontidae, Euteliidae, Erebidae, Nolidae and Noctuidae (Fig. 3). The first two major lineages are well-recognized taxa that have often been considered families within Noctuoidea. Oenosandridae are a small family, only known from Australia, comprising eight species in four genera (Nielsen et al., 1996), which mainly feed on Myrtaceae (Miller, 1991). Notodontidae contain approximately 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 from the rest to warrant full family status. My phylogenetic hypothesis (I) placed previously recognized families—arctiines, lymantriines, aganaines, herminiines and micronoctuines—into the strongly supported Erebidae clade. Within Erebidae, relationships of only a few lineages (e.g., Arctiinae, Aganainae and Herminiinae) were well supported (I). The low support for some nodes within
Erebidae and Noctuidae probably stems from high levels of homoplasy (particularly in the third codon position) and very sparse sampling.
At this stage, with well-supported monophyletic groups established within a phylogenetic framework, the question arose of how best to apply the Linnaean system of nomenclature to the structure of that framework, by making decisions on the content and arrangement of families, subfamilies, tribes and subtribes in a manner 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 well-founded family-level noctuoid classification is certainly an issue of considerable practical importance, because it affects the classification of about a quarter of the world’s lepidopteran species, according to current estimates (Nieukerken et al., 2011). In paper II, I discussed in detail reasons for applying the six family-group system to the higher systematics of Noctuoidea with their advantages and drawbacks.
One of the most striking features of the application of molecular data in my thesis was uncovering the phylogenetic relationships of many unplaced taxa of uncertain affinity. Molecular phylogenetic techniques allowed the easy allocation of these taxa, which had remained ‘incertae sedis’ or with uncertain limits for many years. For example, the position of a number of taxa characterized by the pseudoquadrifine hindwing venation had
Results and Discussion
17
been unstable for a long time (IV–V). Most of these groups (i.e., pseudoquadrifine Noctuidae) were previously assumed to be related to erebid subfamilies (Fibiger & Lafontaine, 2005, Lafontaine & Fibiger, 2006) (II) or even considered 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.
3.2 Pattern of relationships among major lineages of Noctuoidea
Notodontidae are found to be the sister group of all other Noctuoidea, with Oenosandridae branching off next (I). However, this pattern of relationships relative to the rest of Noctuoidea is not well supported in all papers. Both Oenosandridae and Notodontidae have a trifid forewing venation similar to that of Geometridae, a character state that appears to be plesiomorphic relative to the quadrifid forewing venation found in the other 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 and the monophyly of the quadrifid Noctuoidea (i.e., Euteliidae, Erebidae, Nolidae, and Noctuidae). 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) and shared a synapomorphy (i.e., quadrifid forewing venation). The relationships of the 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 in all three phylogenetic methods (MP, ML
and BI). In paper I, Euteliidae were sister to Noctuidae in ML analyses (Fig. 3), and sister to the other three families together in MP analyses. Similarly, in papers IV–V, Euteliidae were placed as sister to the rest of 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 a trichotomy with Erebidae and Noctuidae in MP analyses.
The short internal nodes, with little or no support for many basal divergences, in all quadrifid Noctuoidea lineages (I, II, IV, V), suggest that these groups diversified rapidly within a relatively short period 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 historical explanation (e.g., rapid radiation or the signature of a mass extinction event) or (ii) it is simply an artefact. Under the rapid radiation scenario, it is hypothesized that the lineages diverged so rapidly and within such a narrow time window that there was little opportunity for the ancestors of each monophyletic group to evolve distinctive apomorphies (Futuyma, 2005). However, such a pattern of short internal nodes and lineages with low support can be produced by other factors, such as inadequate data quality, poor and sparse sampling strategy, conflict within or among datasets (data inconsistency), incongruence between the real evolutionary process and the assumed models of sequence evolution, or even lack of phylogenetic signal due to accumulation of overlapping mutations (i.e., the probability of substitutional saturation at a given site).
Results and Discussion
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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 that such phylogenetic patterns may well be signature of mass extinction events (Crisp & Cook, 2009), because mass extinction produces a sharp drop in the cumulative fossil diversity and is commonly thought to stimulate subsequent adaptive radiation,
creating a sharp increase in the rate of diversification (Benton & Emerson, 2007).
Results and Discussion
19
3.3 Phylogenetic hypothesis for quadrifid Noctuoidea
Paper I revealed an urgent need for a comprehensive series of revisions for the higher classifications of quadrifid Noctuoidea families used up to that time. My results in paper I almost failed to recover some previously recognized subfamilies within Erebidae as monophyletic groups. For instance, they suggested that some recent concepts of subfamilies Calpinae, Catocalinae, Erebinae and Phytometrinae were polyphyletic (I). Consequently, the focus in paper II was designed to elucidate the higher-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 each monophyletic lineage (II).
Paper II provides strong support for subordinating five taxa where previously treated as families—Arctiidae, Lymantriidae, Micronoctuidae, Herminiidae and Aganaidae—within Erebidae. One of the most striking features, a strong association among three of them, was shown in papers I–II: Aganainae + Herminiinae + Arctiinae are
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 morphological synapomorphy in the prespiracular position of the counter-tympanal hood, which was until then thought to be plesiomorphic. Adults of many aganaines and arctiines are visually striking and aposematic, and aganaines and herminiines share long labial palps and a bare lower frons. Herminiinae are generally cryptic, feeding on vegetable detritus, and Aganainae are aposematic, feeding on the same suite of toxic cardenolide-synthesizing plant families (Apocynaceae and Moraceae) as the danaine Nymphalidae and other moth genera such as Glyphodes (Crambidae) and Agathia (Geometridae) (Holloway, 2008).
Another interesting result from paper II was the placement of the new established family Micronoctuidae. Hypenodinae are enlarged to incorporate Micronoctuini as a tribe, corroborating the findings of paper I. The subfamily Erebinae, brings together the core catocalines (with the exceptions of Hypocalinae, Toxocampinae and Tinoliinae).
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 in some 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
Results and Discussion
20
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 concept of Calpinae, and supports a monophyletic Calpinae that places members of Anomini and 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 evolved independently (III). The phylogenetic hypothesis suggested three
Results and Discussion
21
subclades for the subfamily Calpinae that were treated as tribes: Phyllodini, Ophiderini and Calpini (II). The polyphyly of the former concept of Calpinae provides an object lesson in how the sharing of peculiar morphological adaptations may mislead classifications, and how shared features of a more subtle nature may be overlooked in an unchallenged traditional classification. Within the entire Lepidoptera, adult hematophagy—the ability to pierce mammalian tissue and extract a blood meal—is limited to species within the vampire moth genus Calyptra Ochsenheimer, which belongs to the subfamily Calpinae. In paper III, we tested whether hematophagy in Calyptra arose from plant (e.g., fruit-piercing) or animal-related behaviors (e.g., tear feeding or lachryphagy). To do that, we subjected the resulting phylogenetic trees to a Bayesian method of ancestral state reconstruction to reconstruct 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 the fruit-piercing habit as opposed to tear feeding or other animal-related feeding behaviors (e.g., dung feeding, urine feeding) (III).
In paper IV, I aimed to elucidate the higher-level phylogeny of Nolidae and to clarify relationships in basal lineages of Noctuidae (V). My phylogenetic hypothesis (I) had already recovered Nolidae and Noctuidae as well-supported monophyletic clades (Fig. 3). The results were fascinating and fairly robust. Many genera previously placed in Nolidae and the former subfamily Ophiderinae (Erebidae) were placed with strong support within Noctuidae, in the subfamily Bagisarinae, supporting an expanded
concept of the subfamily (V). In addition, by increasing taxon sampling of several unassigned Neotropical taxa, I uncovered a previously unknown lineage of Noctuoidea with Neotropical origins (Diphthera Hübner + Lepidodes Guenée) with strong support. This lineage appears to be the sister group to Nolidae, and it shares an unambiguous feature and the most characteristic Nolidae apomorphy yet proposed—the structure of the boat-shaped cocoon with a vertical, anterior exit slit—, suggesting that the clade could be included in the family Nolidae as the subfamily Diphtherinae (IV). Diphtherinae is considered 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 of males, and the presence of a frontal tubercle or process, which is presumably associated with a derived strategy of emergence 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, and Chloephorinae. 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 of whether the hindwings have a trifine or quadrifine venation. To understand character evolution in higher noctuoids and major nolids, I undertook a character optimization analysis based on parsimony
Results and Discussion
22
using the software Mesquite v2.75 (Maddison & Maddison, 2011). My results of ancestral-state reconstructions and character optimization (Fig. 5) showed that most traits—those characters that are considered to be of phylogenetic significance—are clearly synapomorphic for major lineages of quadrifids, except the hindwing venation which was an ambiguous character state (IV). It is generally thought that the pseudoquadrifine 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, and M2 is strong and parallel to M3. My character examinations indicated that the pseudoquadrifine condition is shared by the basal lineages of Noctuidae s.s. (i.e., Bagisarinae, Plusiinae, Dilobinae and Pantheinae) (V), and a few erebid subfamilies (Hypeninae, Herminiinae, Scoliopteryginae and Rivulinae) (II), as well as Diphtherinae (Nolidae) (IV). I evaluated the distribution patterns and evolutionary trends of this complex trait under the two most commonly used character optimization algorithms of parsimony analysis: ACCTRAN—accelerated transformation—and DELTRAN—delayed transformation—(Agnarsson & Miller, 2008). I finally favoured DELTRAN optimization (IV), which minimizes reversals and maximizes convergences and parallel evolution. This favours the acquisition of the derived quadrifine state—base of M2 close to M3—independently in Erebidae, Nolidae (with Diphtherinae excluded), and Euteliidae (Fig. 5). In other words, the plesiomorphic condition of M2 in the hindwing of the four quadrifid lineages is the form found in Diphtherinae and basal lineages of Noctuidae, and arguably some
basal lineages of Erebidae. This trend of character evolution can be traced by checking the position of the vein M2 in different 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 in exactly 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 this condition in some primitive lineages (e.g., Rivulinae, Hypeninae, and Herminiinae). As a consequence, the quadrifine (M2 adjacent to M3) form of venation must have 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 in higher noctuoids. One feature that has been suggested as the prime factor governing the evolution of butterfly-host plant associations, is host growth form, which appears, on the whole, to be more conserved phylogenetically than host-plant taxon 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 and internal nodes indicate the presence of morphological traits as shown below the tree. Names of moths shown in 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 et al., 2006), as it is, apparently, for most
other macrolepidopteran superfamilies (Powell et al., 1998).
Results and Discussion
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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 the plesiomorphic 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—with a worldwide distribution but more diverse in tropics and especially the Neotropics (Weller, 1989, Miller, 1991)—almost all species feed on trees, and only a few are found on herbaceous plants (Miller, 1991). Within Euteliidae, Euteliinae most commonly feed on Anacardiaceae (Powell et al., 1998), a plant family that contains trees and shrubs with highly poisonous flowers. Among their other prominent hosts are Burseraceae (includes both shrubs and trees), Dipterocarpaceae (mainly tropical lowland rainforest trees), Moraceae and Hamamelidaceae (which consists of small trees and shrubs) (Holloway, 1985, Powell et al., 1998). Stictopterinae, the sister group 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, most spectacularly in Arctiinae (Erebidae) and derived trifine lineages (e.g., the pest clade). Although, the great majority of quadrifids feed on living higher plants,
consumption of lower plants and detritus has arisen in several groups, most notably Erebidae (II). For example, lichen-feeding is predominant among lithosiines (Arctiinae) and a number of Aventiini (Boletobiinae) species (Wagner et al., 2008), while detritivory, mycophagy and algivory is dominant in Herminiinae and Boletobiinae (II) and recurs sporadically in other noctuid subfamilies (e.g., some Bryophilinae, see Table 1) (V).
The results in paper V also suggested that Noctuidae included a number of lineages 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 with basal Noctuidae lineages in my phylogenetic hypothesis (Fig. 6)—those characterized by the plesiomorphic pseudoquadrifine condition of the hindwing 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 of major trends in feeding habits in the subfamily Dyopsinae (Table 1) indicates that 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
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Table 1 Larval host-plant families for Noctuidae study taxa. Zoogeographic regions are abbreviated as
follows: P = Palaearctic; O = Oriental; Au = Australasia; Nea = Nearctic; Neo = Neotropical; Af =
Afrotropical.
Results and Discussion
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Continuation of Table 1.
Results and Discussion
27
These results are also widely corroborated by the study of Mitchell et al. (2006), who studied the role of ecological and geographical factors in the diversification of Noctuoidea. They presented a provisional synopsis of species diversity, latitudinal distribution, and host-plant use for major noctuoid groups sampled in their study, superimposed onto the phylogeny. It is revealed that the growth 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 can be reasonably inferred as ancestral for Noctuoidea, as it is, apparently, for most other 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 a general 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 the host-growth form in Noctuoidea is more
evolutionarily conservative than host affiliation (Mitchell et al., 2006). However, my preliminary study of host-plant 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 for studying the evolutionary history of noctuoid moths is to determine the main driving forces behind the diversification of this species-rich group. It has become apparent that the evolution of host-plant use has likely been a key ecological mechanism behind the rapid diversification and evolutionary divergence in the butterfly families (Janz & Nylin, 1998, Janz et al., 2006). Noctuoidea are about three times more diverse than the butterflies, and the role of host plant specialization in the diversification of the moths 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).
Results and Discussion
28
Conclusions and future directions
29
4. CONCLUSIONS AND FUTURE DIRECTIONS
4.1 Conclusions
To conclude my Ph.D. thesis, I have summarized a number of distinctive issues and strategies that I have employed in this thesis that are different from those in previous works.
First of all and probably the most important scheme was the taxon sampling strategy. 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 from significant groupings of genera and morphologically well-supported concepts of higher taxa. In my five projects, I chose up to 393 taxa (Appendix 1) of 45,000 noctuoids that have been formally described, covering almost all recognized major clades of Noctuoidea, as representatives for major lineages of Noctuoidea. 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 morphological traits, in particular, in a species-rich, cosmopolitan and heterogeneous taxon such as Noctuoidea. The high number of species presents complications to the use of morphology and any other kinds of phenotypic traits at this taxonomic level. Morphological data are thus often difficult to homologize and code, require great experience to identify character states
correctly and can be subject to extensive homoplasy (character convergence and reversal). Consequently, despite their major role in inferring phylogenies, morphological analyses have often failed to determine relationships among most groups with confidence. On the other hand, phenotypic traits (e.g., morphological, ecological, host-plant associations, behavioural characters, etc.) and synapomorphies can be properly recognized from molecular phylogeny-based systems. In the context of such a robust evolutionary hypothesis, morphological, ecological and behavioural characters can be better understood. For example, I demonstrated that the prespiracular 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 a common ancestry of these groups within the family Erebidae (II). Polyphyly of the old concept of Calpinae was another clear case (II–III). My results recovered a monophyletic 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 and distant 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 well as its associated modifications of the proboscis and adaptations (III), have
Conclusions and future directions
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evolved independently in Calpinae, Scoliopteryginae (including Anomini), Anobinae and some Erebinae (II). This provides again an object lesson on how the sharing of peculiar morphological adaptations may mislead in classification, and how shared features of a more subtle nature may be overlooked in an unchallenged traditional classification (I). Other examples of discovering morphological apomorphies based on my molecular phylogeny results were in Diphtherinae—characterized by the loss of the proximal pair of tibial spurs on the hindlegs of males and the presence of a frontal tubercle—(IV), Nolidae—construction of a boat-shaped cocoon with a vertical exit slit, and finger-like retinaculum on the forewings of the males—(IV), trifid and quadrifid lineages—condition of vein M2 in forewing—(I, II, IV, V), host-plant associations—e.g., detritivory, lichen-feeding, 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 more derived 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 been previously assigned to various noctuoid groups (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 address a massive and long-recalcitrant phylogenetic problem, that of the relationships among and within major lineages of the largest superfamily of Lepidoptera, Noctuoidea. The phylogenetic analyses that I employed were well designed, and many relatively deep nodes are strongly resolved and substantial progress has been made on the backbone phylogeny of Noctuoidea. This group of insects is of major economic and ecological importance. It constitutes an exemplar case in which molecular methods, which seem to be highly informative at this level, have been of enormous 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 the evolution of Noctuoidea, but there are still many unanswered questions. For example, it is crucial to determine where, when and how the major groups of noctuoids diverged and evolved. The common factors that influence the speciation process and identifying possible reasons for this remarkable and extraordinary diversity of species among other herbivorous insects are not yet understood. Likewise, the patterns of diversification, the main driving forces behind the diversification of this species-rich group, and plausible explanations for the differences in diversity among the various groups within Noctuoidea have yet to be determined—for instance, why does a family such as Euteliidae contain about 500 species, but another like Erebidae contains 25,000 species? It would be of great interest to know whether major climatic changes and mass extinction events over geological
Conclusions and future directions
31
time scales have had a major impact on the diversification of Noctuoidea. And it would be most interested to infer a scenario of the evolutionary history and biogeography of noctuoids, based on all available data (i.e., morphology, DNA sequences, ecological data, geographical distributions, and geological and paleontological information).
Further studies are also needed to identify the reasons and causes for the short basal branches, i.e., whether there is a historical explanation behind them (e.g., rapid radiation), or whether it is simply an artefact of insufficient data.
The use of novel tools in DNA sequencing technologies, such as Next-Generation Sequencing (NGS) methods and relatively new field of phylogenomics, might be able to address the state of uncertainty in Noctuoidea diversification and their rapid radiations. Phylogenomics is useful for evolutionary studies, in particular resolving ambiguous phylogenies and for verifying relationships created on the basis of a few gene regions (Hackett et al., 2008). Since the cost of whole genome sequencing is decreasing, anticipation of total genome sequences from the major lineages of interest, is no longer a distant dream (Murphy et al., 2004).
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5. ACKNOWLEDGEMENTS
Laboratory work for this project was funded through a grant from the Academy of Finland (grant no. 129811) and Kone Foundation to my supervisor, Dr. Niklas Wahlberg. My subsistence expenses were funded by the CIMO (May 2008 to April 2009), the Finnish Cultural Foundation (February 2010 to January 2011) and the Alfred Kordelin Foundation (January 2012 to June 2012). I would like to take this opportunity 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 first day I started my laboratory work (in September 2008) to the final parts, where I am typing these words (April 2012). I have started from below zero when Niklas taught me how to work in Genetics laboratory! It was a completely 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 and confidence you had on me.
I would like to dedicate this thesis to one of the most influential persons in my life, the late Michael 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 made this thesis possible. I think a nice feature of this thesis was the inclusion of three leading authorities on noctuoid morphology and classification in the world, and that they have been closely involved in the design and interpretation 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 your confidence in me. Ian and Jeremy, I would like to thank you for your fruitful, brilliant and comprehensive comments on the manuscripts.
I am pleased to acknowledge my co-authors, Dr. Lauri Kaila and Dr. Marko Mutanen, who have greatly contributed to the improvement of manuscripts and also for sequencing of some material. I sincerely thank to 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 and valuable comments on our joint-papers (IV,V). I also wish to acknowledge the work of Ms Jocelyn Gill (CNC) for her expert work on the color plates for paper IV.
A large number of specimens for the molecular study were provided by a group of nice people collaborating with us in this project: 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 and Jaakko 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
33
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 national d'Histoire naturelle, France); Donald Lafontaine and Christian Schmidt (the Canadian National Collection of Insects); David Wagner (University of Connecticut, USA); Vasiliy Kravchenko (Tel Aviv University, Israel), as well as a number of private 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 the thesis, Prof. Charlie Mitter and Dr. Tommi Nyman, for their informative comments on the thesis introduction. Despite not having had the chance to meet Prof. Mitter in person, I have learned a lot from his great personality. I was impressed, for the first time in October 2007, before coming to Finland, when we were asking 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 my doctoral 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 technical support.
My sincere thank goes to Dr. Carlos Peña, NSG’ web application developer, who created the 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 nice moments we have shared together in the same office and coffee room. Kalle, thank you so much for being positive, cheerful, joyful and
helpful all the time. I will never forget our Finnish sauna events, will never forget Vappu 2010 and 2011! I would like to thank Julien Leneveu and Heike Witthauer for providing me with the great atmosphere in which to work in the TEGlab environment. I also wish to thank Raija Rouhiainen, who was the one who took care of my financial matters, calculating Niklas’s grants several times a year.
I would like to thank Heidi and Johanna for keeping 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 the lecturers 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 grateful to all the TEGlab people, in particular: Tatjaana, Mikhail, Roghelio, Paula, Veronika, Elsi, Heidi, Megha, Ksenia, Walter, Bineet and Eero for technical help, resolving practical issues, and of course tolerating me for four years! Also, it is my privilege to share my feelings of gratitude with my always cheerful friends, 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 and desperate.
And finally thanks to Suomi for letting me be here for four years and allowing me finally to live! I have experienced things I have never experienced 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!!!
References
34
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Appendix
37
APPENDIX F
amily
Subf
amily
Spec
ies
Pap
er
Specimen ID
COI-LCO
COI-Jerry
EF1
-egin
EF1
-en
Wingless
GAPDH
RpS5
MDH
CAD
IDH
Type status
Locality
Out
grou
p
Dre
pan
idae
Thy
atir
inae
Thy
atir
a ba
tis
IM
M00
027
GU
8285
80G
U82
8380
GU
8289
19G
U82
9212
GU
8294
81G
U82
9743
GU
8305
97G
U83
0293
GU
8280
83G
U82
9969
TG
/TS
FIN
LA
ND
Sp
hing
idae
Sphi
ngin
aeSp
hinx
ligu
stri
IN
W14
1-12
EU
1413
58E
U14
1358
EU
1366
65E
U13
6665
EU
1412
39E
U14
1494
EU
1413
91E
U14
1615
EU
1413
13E
U14
1550
TG
/TS
FIN
LA
ND
Bom
byc
idae
Bom
byci
nae
Bom
byx
mor
iI
NW
149-
1E
U14
1360
EU
1413
60E
U13
6667
EU
1366
67E
U14
1241
EU
1414
95E
U14
1393
EU
1416
17E
U14
1315
EU
1415
52T
G/T
SU
SA
Geo
met
rid
aeA
rchi
eari
nae
Arc
hiea
ris
part
heni
asI
NW
107-
1D
Q01
8928
DQ
0189
28D
Q01
8899
DQ
0188
99D
Q01
8869
EU
1414
85E
U14
1381
EU
1416
04E
U14
1303
EU
1415
39T
G/T
SS
WE
DE
N
Ingr
oup
Oen
osan
drid
aeO
enos
andr
a bo
isdu
vali
I,II
,IV
MM
0759
0G
U82
8791
GU
9297
62G
U82
9098
GU
8293
77G
U82
9651
GU
8298
71G
U83
0751
GU
8304
92G
U82
8266
GU
8301
73T
G/T
SA
US
TR
AL
IA
Oen
osan
drid
aeD
isco
phle
bia
sp.
I,IV
RZ
403
HQ
0062
17H
Q00
6921
HQ
0063
13H
Q00
6404
HQ
0068
25H
Q00
6480
HQ
0067
29H
Q00
6638
−H
Q00
6551
AU
ST
RA
LIA
Not
odon
tida
ePh
aler
inae
Pha
lera
buc
epha
laI,
IIM
M00
122
GU
8286
07G
U82
8405
GU
8289
41G
U82
9235
GU
8295
02−
GU
8306
17G
U83
0318
GU
8281
08G
U82
9995
TG
/TS
FIN
LA
ND
Not
odon
tida
eH
eter
ocam
pina
eSt
auro
pus
fagi
I,II
,IV
MM
0098
1G
U82
8651
GU
8284
49G
U82
8983
GU
8292
66G
U82
9539
GU
8297
80G
U83
0650
GU
8303
57G
U82
8148
GU
8300
38T
SFI
NL
AN
D
Not
odon
tida
eN
otod
ontin
aeN
otod
onta
dro
med
ariu
sI,
II,I
V,V
MM
0099
8G
U82
8653
GU
8284
51G
U82
8984
GU
8292
68G
U82
9540
GU
8297
81G
U83
0652
GU
8303
59G
U82
8150
GU
8300
40T
G/T
SFI
NL
AN
D
Not
odon
tida
ePy
gaer
inae
Clo
ster
a pi
gra
IM
M01
005
GU
8286
54G
U82
8452
GU
8289
85G
U82
9269
GU
8295
41G
U82
9782
GU
8306
53G
U83
0360
GU
8281
51G
U83
0041
FIN
LA
ND
Not
odon
tida
eT
haum
etop
oein
aeE
pico
ma
mel
anos
tict
aI
MM
0759
2G
U82
8792
GU
9297
63G
U82
9099
GU
8293
78G
U82
9652
GU
8298
72G
U83
0752
GU
8304
93G
U82
8267
GU
8301
74A
US
TR
AL
IA
Not
odon
tida
eT
haum
etop
oein
aeT
haum
etop
oea
soli
tari
aI,
VM
M09
888
GU
8288
43G
U92
9807
GU
8291
44−
GU
8296
92G
U82
9904
GU
8307
91G
U83
0534
GU
8283
07G
U83
0223
TG
GR
EE
CE
Not
odon
tida
eD
udus
inae
Cri
node
s be
scke
iI
05-S
RN
P-5
7213
GU
8285
27−
GU
8288
73G
U82
9175
GU
8294
34−
GU
8305
63G
U83
0251
GU
8280
39G
U82
9918
CO
ST
A R
ICA
Not
odon
tida
eN
ysta
lein
aeN
ysta
lea
stri
ata
I05
-SR
NP
-444
3G
U82
8525
−G
U82
8871
GU
8291
73G
U82
9432
GU
8297
17G
U83
0561
GU
8302
49G
U82
8037
GU
8299
16T
GC
OS
TA
RIC
A
Not
odon
tida
eD
iopt
inae
Scot
ura
leuc
ophl
eps
I06
-SR
NP
-227
81G
U82
8532
GU
8283
34G
U82
8878
GU
8291
79G
U82
9439
GU
8297
21G
U83
0568
GU
8302
56G
U82
8044
GU
8299
23C
OS
TA
RIC
A
Eut
eliid
aeE
utel
iinae
Eut
elia
adu
latr
ixI,
II,V
MM
0016
0G
U82
8621
GU
8284
19G
U82
8956
GU
8292
46G
U82
9516
GU
8297
64G
U83
0629
GU
8303
30G
U82
8122
GU
8300
10T
G/T
SG
RE
EC
E
Eut
eliid
aeE
utel
iinae
Eut
elia
gey
eri
IVR
Z50
8X
XX
XX
XX
X−
XT
GH
ON
G K
ON
G
Eut
eliid
aeE
utel
iinae
Mar
athy
ssa
basa
lis
IR
Z23
HQ
0061
83H
Q00
6887
HQ
0062
79H
Q00
6374
HQ
0067
91H
Q00
6455
HQ
0066
98H
Q00
6606
HQ
0069
79H
Q00
6528
TS
US
A
Eut
eliid
aeE
utel
iinae
Tar
gall
a su
boce
llat
aI,
II,I
V,V
RZ
35H
Q00
6210
HQ
0069
14H
Q00
6306
HQ
0063
97H
Q00
6818
HQ
0064
73H
Q00
6722
HQ
0066
31H
Q00
7000
−H
ON
G K
ON
G
Eut
eliid
aeSt
icto
pter
inae
Aeg
ilia
des
crib
ens
IIR
Z28
7JN
4012
34JN
4011
18JN
4013
52−
JN40
0927
JN40
1566
JN40
1873
JN40
1772
JN40
1036
−T
SIN
DO
NE
SIA
Eut
eliid
aeSt
icto
pter
inae
Lop
hopt
era
hem
ithy
ris
I,II
MM
0761
4G
U82
8802
GU
9297
72G
U82
9107
GU
8293
85G
U82
9661
GU
8298
79G
U83
0759
GU
8305
01G
U82
8274
GU
8301
83A
US
TR
AL
IA
Eut
eliid
aeSt
icto
pter
inae
Lop
hopt
era
squa
mm
iger
aIV
,VR
Z12
0X
XX
XX
XX
X−
XT
GH
ON
G K
ON
G
Eut
eliid
aeSt
icto
pter
inae
Stic
topt
era
colu
mba
IV,V
RZ
541
XX
XX
X−
XX
−X
TG
MA
LA
YS
IA
Ere
bida
eSc
olio
pter
ygin
aeSc
olio
pter
yx li
batr
ixI,
II,I
VM
M00
407
GU
8286
41G
U82
8439
GU
8289
75G
U82
9260
GU
8295
32−
GU
8306
43G
U83
0348
GU
8281
40G
U83
0028
TG
/TS
FIN
LA
ND
Ere
bida
eSc
olio
pter
ygin
aeO
sson
oba
torp
ida
IIR
Z41
1JN
4012
52JN
4011
34JN
4013
69JN
4014
80JN
4009
39JN
4015
82JN
4018
93−
JN40
1050
−T
SM
AL
AY
SIA
Ere
bida
eSc
olio
pter
ygin
aeR
usic
ada
fulv
ida
IIR
Z10
1JN
4012
53JN
4011
35JN
4013
70JN
4014
81JN
4009
83JN
4015
83JN
4018
94JN
4017
90JN
4010
51JN
4016
83H
ON
G K
ON
G
Ere
bida
eSc
olio
pter
ygin
aeR
usic
ada
met
axan
tha
I,II
RZ
55H
Q00
6227
HQ
0069
30H
Q00
6322
HQ
0064
14H
Q00
6835
−H
Q00
6739
HQ
0066
47H
Q00
7016
HQ
0065
60T
GT
AIW
AN
Ere
bida
eSc
olio
pter
ygin
aeG
onit
is in
volu
taI,
IIR
Z13
HQ
0061
66H
Q00
6963
HQ
0062
63H
Q00
6357
HQ
0067
75−
HQ
0066
82H
Q00
6592
HQ
0069
63−
TG
TA
NZ
AN
IA
Ere
bida
eSc
olio
pter
ygin
aeA
nom
is fl
ava
IIR
Z10
0JN
4012
54JN
4011
36JN
4013
71JN
4014
82JN
4009
81JN
4015
84−
−JN
4010
52JN
4016
84H
ON
G K
ON
G
Ere
bida
eun
assi
gned
Rhe
sala
impa
rata
IIR
Z26
5JN
4012
55JN
4011
37JN
4013
72JN
4014
83JN
4009
40JN
4015
85−
JN40
1791
JN40
1053
JN40
1685
TS
HO
NG
KO
NG
Ere
bida
eun
assi
gned
Nyc
hiop
tera
noct
uida
lis
IIR
Z28
3JN
4012
56JN
4011
38JN
4013
73−
JN40
0941
−−
JN40
1792
−−
TS
US
A
Ere
bida
eR
ivul
inae
Riv
ula
seri
ceal
isI,
IIM
M01
404
GU
8286
64G
U82
8462
GU
8289
95G
U82
9278
−G
U82
9791
−G
U83
0370
GU
8281
61G
U83
0051
TG
/TS
FIN
LA
ND
Appendix
38
Ere
bida
eR
ivul
inae
Riv
ula
ochr
eaII
RZ
159
JN40
1257
JN40
1139
JN40
1374
JN40
1484
JN40
0979
JN40
1586
−JN
4017
93JN
4010
54−
TG
GH
AN
A
Ere
bida
eR
ivul
inae
Oxy
cill
a on
doI,
IIR
Z24
HQ
0061
84H
Q00
6888
HQ
0062
80H
Q00
6375
HQ
0067
92H
Q00
6456
−H
Q00
6607
HQ
0069
80H
Q00
6529
US
A
Ere
bida
eR
ivul
inae
Boc
ula
bifa
ria
IIR
Z41
3JN
4012
58JN
4011
40JN
4013
75JN
4014
85JN
4009
42JN
4015
87−
−JN
4010
55JN
4016
86M
AL
AY
SIA
Ere
bida
eR
ivul
inae
Ogl
asa
anso
rgei
IIR
Z16
7JN
4012
59JN
4011
41JN
4013
76JN
4014
86JN
4009
86JN
4015
88−
−JN
4010
56JN
4016
87G
HA
NA
Ere
bida
eR
ivul
inae
Ale
sua
etia
lis
IIR
Z94
JN40
1260
JN40
1142
JN40
1377
JN40
1487
JN40
0943
JN40
1589
−JN
4017
94−
JN40
1688
TS
CO
ST
A R
ICA
Ere
bida
eA
nobi
nae
Ano
ba a
ngul
ipla
gaI,
IIR
Z33
2H
Q00
6206
HQ
0069
10H
Q00
6302
HQ
0063
95H
Q00
6814
HQ
0064
69−
HQ
0066
27−
HQ
0065
44T
GG
HA
NA
Ere
bida
eA
nobi
nae
Mar
cipa
sp.
I,II
RZ
177
HQ
0061
77H
Q00
6881
−H
Q00
6368
HQ
0067
85H
Q00
6450
−H
Q00
6601
HQ
0069
73H
Q00
6522
GH
AN
A
Ere
bida
eA
nobi
nae
Mar
cipa
sp.
IIR
Z20
0JN
4012
61JN
4011
43JN
4013
78JN
4014
88JN
4009
44JN
4015
90−
JN40
1795
−JN
4016
89G
HA
NA
Ere
bida
eA
nobi
nae
Ple
copt
era
maj
orII
RZ
183
JN40
1262
JN40
1144
−JN
4014
89JN
4009
45JN
4015
91−
JN40
1796
−JN
4016
90G
HA
NA
Ere
bida
eA
nobi
nae
Cri
thot
e pr
omin
ens
IIR
Z10
9JN
4012
63JN
4011
45JN
4013
79JN
4014
90JN
4009
46JN
4015
92−
JN40
1797
JN40
1057
JN40
1691
HO
NG
KO
NG
Ere
bida
eA
nobi
nae
Rem
a co
stim
acul
aII
RZ
103
JN40
1264
JN40
1146
JN40
1380
JN40
1491
JN40
0947
JN40
1593
−JN
4017
98JN
4010
58JN
4016
92T
SH
ON
G K
ON
G
Ere
bida
eA
nobi
nae
Ban
iana
str
igat
aII
RZ
92JN
4012
65JN
4011
47JN
4013
81JN
4014
92JN
4009
48JN
4015
94−
JN40
1799
JN40
1059
JN40
1693
CO
ST
A R
ICA
Ere
bida
eA
nobi
nae
Dei
nopa
sig
nipl
ena
IIR
Z31
1JN
4012
66JN
4011
48JN
4013
82JN
4014
93JN
4009
49JN
4015
95−
JN40
1800
JN40
1060
JN40
1694
CO
ST
A R
ICA
Ere
bida
eH
ypen
inae
Hyp
ena
prob
osci
dali
sI,
IIM
M01
545
GU
8286
68G
U82
8466
GU
8289
99G
U82
9282
GU
8295
53G
U82
9794
GU
8306
64G
U83
0374
GU
8281
65G
U83
0055
TG
/TS
FIN
LA
ND
Ere
bida
eH
ypen
inae
Hyp
ena
balt
imor
alis
IIR
Z36
7JN
4012
67JN
4011
49JN
4013
83JN
4014
94JN
4009
93JN
4015
96JN
4018
95JN
4018
01JN
4010
61JN
4016
95T
GU
SA
Ere
bida
eH
ypen
inae
Hyp
ena
lace
rata
lis
IIR
Z36
8JN
4012
68JN
4011
50JN
4013
84JN
4014
95JN
4009
50−
JN40
1896
−JN
4010
62JN
4016
96T
GH
ON
G K
ON
G
Ere
bida
eun
assi
gned
Cul
trip
alpa
sp.
IIR
Z39
4JN
4012
69JN
4011
51JN
4013
85JN
4014
96JN
4009
51JN
4015
97JN
4018
97JN
4018
02−
JN40
1697
MA
LA
YS
IA
Ere
bida
eun
assi
gned
Col
oboc
hyla
sal
ical
isI,
IIR
Z4
HQ
0062
15H
Q00
6919
HQ
0063
11H
Q00
6402
HQ
0068
23H
Q00
6478
HQ
0067
27H
Q00
6636
HQ
0070
05−
TS
HU
NG
AR
Y
Ere
bida
eL
yman
triin
aeL
yman
tria
mon
acha
I,II
,IV
,VM
M01
048
GU
8286
55G
U82
8453
GU
8289
86G
U82
9270
GU
8295
42−
GU
8306
54G
U83
0361
GU
8281
52G
U83
0042
TG
FIN
LA
ND
Ere
bida
eL
yman
triin
aeL
euco
ma
sali
cis
I,II
MM
0674
0G
U82
8748
GU
9297
22G
U82
9062
GU
8293
47G
U82
9611
−G
U83
0719
GU
8304
49G
U82
8232
GU
8301
32T
G/T
SFI
NL
AN
D
Ere
bida
eL
yman
triin
aeN
ygm
ia p
lana
I,II
RZ
34H
Q00
6209
HQ
0069
13H
Q00
6305
HQ
0063
96H
Q00
6817
HQ
0064
72H
Q00
6721
HQ
0066
30H
Q00
6999
HQ
0065
46T
GH
ON
G K
ON
G
Ere
bida
eL
yman
triin
aeO
rgyi
a an
tiqu
aI,
IIR
Z13
0H
Q00
6167
HQ
0069
64H
Q00
6264
HQ
0063
58H
Q00
6776
HQ
0064
43H
Q00
6683
HQ
0065
93H
Q00
6964
HQ
0065
13T
G/T
SFI
NL
AN
D
Ere
bida
eL
yman
triin
aeA
rcto
rnis
sp.
I,II
RZ
89H
Q00
6241
HQ
0069
43H
Q00
6335
HQ
0064
28H
Q00
6849
HQ
0064
94H
Q00
6752
HQ
0066
59H
Q00
7024
HQ
0065
72T
GJA
PA
N
Ere
bida
ePa
ngra
ptin
aeP
angr
apta
bic
ornu
taI,
IIR
Z40
HQ
0062
16H
Q00
6920
HQ
0063
12H
Q00
6403
HQ
0068
24H
Q00
6479
HQ
0067
28H
Q00
6637
HQ
0070
06H
Q00
6550
TG
HO
NG
KO
NG
Ere
bida
ePa
ngra
ptin
aeP
angr
apta
dec
oral
isI,
IIR
Z66
HQ
0062
36H
Q00
6939
HQ
0063
31H
Q00
6423
HQ
0068
44−
HQ
0067
47−
HQ
0070
22H
Q00
6568
TG
/TS
US
A
Ere
bida
ePa
ngra
ptin
aeC
hrys
ogra
pta
igne
ola
IIR
Z40
8JN
4012
70JN
4011
52JN
4013
86JN
4014
97JN
4009
52JN
4015
98JN
4018
98JN
4018
03−
−T
SM
AL
AY
SIA
Ere
bida
ePa
ngra
ptin
aeH
ypos
eman
sis
sing
haII
RZ
279
JN40
1271
JN40
1153
JN40
1387
JN40
1498
JN40
0953
JN40
1599
JN40
1899
JN40
1804
JN40
1063
JN40
1698
TS
MA
LA
YS
IA
Ere
bida
ePa
ngra
ptin
aeG
raci
lode
s ca
ffra
IIR
Z29
2JN
4012
72JN
4011
54JN
4013
88JN
4014
99JN
4009
54JN
4016
00JN
4019
00JN
4018
05JN
4010
64JN
4016
99T
ST
AN
ZA
NIA
Ere
bida
ePa
ngra
ptin
aeE
pisp
aris
cos
tist
riga
IIR
Z31
9JN
4012
73JN
4011
55JN
4013
89JN
4015
00JN
4009
55JN
4016
01JN
4019
01−
−JN
4017
00M
AL
AY
SIA
Ere
bida
ePa
ngra
ptin
aeM
asca
aba
ctal
isI,
IIR
Z18
HQ
0061
78H
Q00
6882
HQ
0062
74H
Q00
6369
HQ
0067
86H
Q00
6451
HQ
0066
93−
HQ
0069
74H
Q00
6523
TS
IND
ON
ES
IA
Ere
bida
eun
assi
gned
Schi
stor
hynx
arge
ntis
trig
aII
RZ
119
JN40
1274
JN40
1156
JN40
1390
JN40
1501
JN40
0956
JN40
1602
JN40
1902
JN40
1806
JN40
1065
JN40
1701
TS
HO
NG
KO
NG
Ere
bida
eH
erm
iniin
aeP
olyp
ogon
str
igil
atus
I,II
MM
0128
6G
U82
8663
GU
8284
61G
U82
8994
GU
8292
77G
U82
9549
GU
8297
90G
U83
0660
GU
8303
69G
U82
8160
GU
8300
50T
G/T
SFI
NL
AN
D
Ere
bida
eH
erm
iniin
aeP
arac
olax
tris
talis
I,II
RZ
5H
Q00
6224
HQ
0069
27H
Q00
6319
HQ
0064
11H
Q00
6832
−H
Q00
6736
−H
Q00
7013
−H
UN
GA
RY
Ere
bida
eH
erm
iniin
aeH
erm
inia
tars
icri
nali
sI,
IIR
Z6
HQ
0062
32H
Q00
6935
HQ
0063
27H
Q00
6419
HQ
0068
40H
Q00
6489
−−
−−
TG
HU
NG
AR
Y
Ere
bida
eH
erm
iniin
aeSi
mpl
icia
sp.
I,II
RZ
166
HQ
0061
75H
Q00
6879
HQ
0062
72H
Q00
6366
−H
Q00
6448
HQ
0066
91H
Q00
6599
HQ
0069
71H
Q00
6520
GH
AN
A
Ere
bida
eH
erm
iniin
aeId
ia a
emul
aII
RZ
271
JN40
1275
JN40
1157
JN40
1391
JN40
1502
JN40
0957
−−
JN40
1807
JN40
1066
JN40
1702
TS
US
A
Ere
bida
eH
erm
iniin
aeL
ysim
elia
nel
eusa
lis
IIR
Z26
0JN
4012
76JN
4011
58−
JN40
1503
JN40
0958
JN40
1603
−JN
4018
08JN
4010
67JN
4017
03T
SH
ON
G K
ON
G
Ere
bida
eH
erm
iniin
aeN
odar
ia v
erti
cali
sII
RZ
180
JN40
1277
JN40
1159
JN40
1392
JN40
1504
JN40
0959
JN40
1604
−−
−JN
4017
04G
HA
NA
Appendix
39
Ere
bida
eA
gana
inae
Aso
ta c
aric
aeI,
IIM
M00
145
GU
8286
15G
U82
8413
GU
8289
49G
U82
9240
GU
8295
09−
GU
8306
24G
U83
0325
GU
8281
15G
U83
0003
TG
/TS
TA
HIL
AN
D
Ere
bida
eA
gana
inae
Aso
ta h
elic
onia
I,II
RZ
44H
Q00
6220
HQ
0069
24H
Q00
6316
HQ
0064
07H
Q00
6828
HQ
0064
83H
Q00
6732
HQ
0066
41H
Q00
7009
HQ
0065
54T
GH
ON
G K
ON
G
Ere
bida
eA
gana
inae
Neo
cher
a in
ops
IIR
Z34
6JN
4012
78JN
4011
60JN
4013
93−
JN40
0960
JN40
1605
−JN
4018
09JN
4010
68JN
4017
05JA
PA
N
Ere
bida
eA
gana
inae
Eup
loci
a m
embl
iari
aII
RZ
345
JN40
1279
JN40
1161
JN40
1394
JN40
1505
JN40
0961
JN40
1606
−JN
4018
10−
JN40
1706
TS
MA
LA
YS
IA
Ere
bida
eA
gana
inae
Per
idro
me
orbi
cula
ris
IIR
Z28
0JN
4012
80JN
4011
62JN
4013
95JN
4015
06JN
4009
62JN
4016
07JN
4019
03JN
4018
11−
JN40
1707
TS
MA
LA
YS
IA
Ere
bida
eA
gana
inae
Mec
odin
a pr
aeci
pua
IIR
Z26
8JN
4012
81JN
4011
63JN
4013
96JN
4015
07JN
4009
63JN
4016
08−
JN40
1812
JN40
1069
JN40
1708
HO
NG
KO
NG
Ere
bida
eA
gana
inae
Psi
mad
a qu
adri
penn
isII
RZ
47JN
4012
82JN
4011
64JN
4013
97JN
4015
08JN
4009
64−
JN40
1904
JN40
1813
JN40
1070
JN40
1709
TG
/TS
HO
NG
KO
NG
Ere
bida
eA
rctii
nae
Bru
nia
anti
caI,
IIR
Z28
HQ
0061
93H
Q00
6897
HQ
0062
89H
Q00
6383
HQ
0068
01H
Q00
6462
HQ
0067
06H
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6614
−H
Q00
6534
TS
HO
NG
KO
NG
Ere
bida
eA
rctii
nae
Gar
udin
ia s
imul
ana
IIR
Z39
9JN
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83JN
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65JN
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98JN
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09JN
4009
65JN
4016
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05JN
4018
14JN
4010
71JN
4017
10M
AL
AY
SIA
Ere
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eA
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nae
Eug
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RZ
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JN40
1284
JN40
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06JN
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11M
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Ere
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Cya
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RZ
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JN40
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JN40
1816
JN40
1073
JN40
1712
MA
LA
YS
IA
Ere
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eA
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nae
Bar
sine
sp
IIR
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7JN
4012
86JN
4011
68JN
4014
01JN
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12JN
4009
68−
JN40
1878
JN40
1817
JN40
1074
JN40
1713
MA
LA
YS
IA
Ere
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eA
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nae
Api
sa c
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cens
I,II
MM
0584
3H
Q00
6146
HQ
0068
53−
HQ
0063
39H
Q00
6765
−H
Q00
6663
−−
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SO
MA
N
Ere
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nae
Synt
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phe
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RZ
8H
Q00
6238
HQ
0069
41−
HQ
0064
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Q00
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HQ
0064
92H
Q00
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HQ
0066
56−
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G/T
SH
UN
GA
RY
Ere
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nae
Dys
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mul
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M00
154
GU
8286
19G
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GU
8289
54G
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9244
GU
8295
14−
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GU
8281
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0008
GR
EE
CE
Ere
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nae
Ant
ichl
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vir
idis
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MM
0538
0H
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HQ
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79H
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Ere
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8288
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8294
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8305
70−
−G
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9926
TG
US
A
Ere
bida
eA
rctii
nae
Cos
cini
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I,II
MM
0567
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HQ
0068
56H
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6247
HQ
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HQ
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Ere
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GU
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80G
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9180
GU
8294
41−
GU
8305
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0258
GU
8280
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9925
CO
ST
A R
ICA
Ere
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nae
Dys
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HQ
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HQ
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27H
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6848
HQ
0064
93H
Q00
6751
HQ
0066
58−
HQ
0065
71C
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TA
RIC
A
Ere
bida
eA
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nae
Nyc
tem
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baul
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RZ
387
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1287
JN40
1169
JN40
1402
JN40
1513
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0969
JN40
1611
JN40
1909
JN40
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4017
14M
AL
AY
SIA
Ere
bida
eA
rctii
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Cal
lim
orph
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min
ula
I,II
RZ
136
HQ
0061
69H
Q00
6873
HQ
0062
66H
Q00
6360
HQ
0067
78H
Q00
6444
HQ
0066
85H
Q00
6594
HQ
0069
65H
Q00
6514
TG
/TS
RU
SS
IA
Ere
bida
eA
rctii
nae
Cre
aton
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sien
sI,
IIR
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HQ
0061
98H
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6902
HQ
0062
94H
Q00
6387
HQ
0068
06−
HQ
0067
11H
Q00
6619
HQ
0069
91H
Q00
6537
HO
NG
KO
NG
Ere
bida
eA
rctii
nae
Arc
tia c
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I,II
,IV
,VM
M03
713
GU
8286
93G
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8489
−G
U82
9305
GU
8295
73G
U82
9813
−G
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0398
GU
8281
85G
U83
0080
TG
/TS
FIN
LA
ND
Ere
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eA
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nae
Am
eril
a as
treu
sII
RZ
404
JN40
1288
JN40
1170
JN40
1403
JN40
1514
−JN
4016
12JN
4019
10−
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4017
15T
SM
AL
AY
SIA
Ere
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RZ
93JN
4012
89JN
4011
71JN
4014
04JN
4015
15JN
4009
70JN
4016
13JN
4019
11JN
4018
19−
JN40
1716
CO
ST
A R
ICA
Ere
bida
eC
alpi
nae
Phy
llod
es e
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ovii
I,II
,III
RZ
56H
Q00
6228
HQ
0069
31H
Q00
6323
HQ
0064
15H
Q00
6836
−H
Q00
6740
HQ
0066
48−
HQ
0065
61T
GT
AIW
AN
Ere
bida
eC
alpi
nae
Phy
llod
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peri
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II,I
IIR
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6JN
6748
69JN
6748
51JN
6748
86JN
6749
02JN
6749
68JN
6749
19JN
6749
50JN
6749
34−
JN67
4991
TG
AU
ST
RA
LIA
Ere
bida
eC
alpi
nae
Min
iode
s ph
aeos
oma
I,II
,III
RZ
153
HQ
0061
73H
Q00
6877
HQ
0062
70H
Q00
6364
HQ
0067
82H
Q00
6446
HQ
0066
89H
Q00
6597
HQ
0069
69H
Q00
6518
GH
AN
A
Ere
bida
eC
alpi
nae
Hem
icer
atoi
des
sitt
aca
II,I
IIR
Z15
5JN
4012
90JN
4011
72JN
4014
05JN
4015
16JN
4009
71JN
4016
14JN
4019
12JN
4018
20−
JN40
1717
GH
AN
A
Ere
bida
eC
alpi
nae
Eud
ocim
a sa
lam
inia
II,I
IIR
Z33
8JN
4012
91JN
4011
73JN
4014
06JN
4015
17JN
4009
90JN
4016
15JN
4019
13JN
4018
21−
JN40
1740
TG
/TS
HO
NG
KO
NG
Ere
bida
eC
alpi
nae
Eud
ocim
a fu
llon
iaI,
II,I
IIR
Z16
HQ
0061
74H
Q00
6878
HQ
0062
71H
Q00
6365
HQ
0067
83H
Q00
6447
HQ
0066
90H
Q00
6598
HQ
0069
70H
Q00
6519
TG
MA
LA
YS
IA
Ere
bida
eC
alpi
nae
Eud
ocim
a di
viti
osa
III
RZ
210
JN67
4870
JN67
4852
JN67
4887
JN67
4903
JN67
4969
JN67
4920
JN67
4951
−−
JN67
4992
TG
GH
AN
A
Ere
bida
eC
alpi
nae
Eud
ocim
a ty
rann
usII
IR
Z43
0JN
6748
71JN
6748
53JN
6748
88JN
6749
04JN
6749
70JN
6749
21JN
6749
52JN
6749
35−
JN67
4993
TG
RU
SS
IA
Ere
bida
eC
alpi
nae
Gon
odon
ta u
xor
I,II
,III
RZ
335
HQ
0062
08H
Q00
6912
HQ
0063
04−
HQ
0068
16H
Q00
6471
HQ
0067
20H
Q00
6629
−H
Q00
6545
CO
ST
A R
ICA
Ere
bida
eC
alpi
nae
Gon
odon
ta li
ncus
III
RZ
417
JN67
4872
JN67
4854
JN67
4889
JN67
4905
JN67
4971
JN67
4922
JN67
4953
JN67
4936
−JN
6749
94B
RA
ZIL
Ere
bida
eC
alpi
nae
Gon
odon
ta m
illa
III
RZ
421
JN67
4873
JN67
4855
JN67
4890
JN67
4906
JN67
4972
JN67
4923
JN67
4954
JN67
4937
−JN
6749
95B
RA
ZIL
Ere
bida
eC
alpi
nae
Gon
odon
ta s
yrna
III
RZ
420
JN67
4874
JN67
4856
JN67
4891
JN67
4907
JN67
4973
JN67
4924
JN67
4955
JN67
4938
−JN
6749
96B
RA
ZIL
Appendix
40
Ere
bida
eC
alpi
nae
Gon
odon
ta fu
lvan
gula
III
RZ
423
JN67
4875
JN67
4857
JN67
4892
JN67
4908
JN67
4974
−JN
6749
57JN
6749
40−
JN67
4997
BR
AZ
IL
Ere
bida
eC
alpi
nae
Gon
odon
ta n
utri
xII
IR
Z43
2JN
6748
76JN
6748
58JN
6748
93JN
6749
09JN
6749
75−
JN67
4956
JN67
4939
JN67
4985
JN67
4998
US
A
Ere
bida
eC
alpi
nae
Gon
odon
ta s
iche
asII
IR
Z41
9JN
6748
77JN
6748
59JN
6748
94JN
6749
10JN
6749
76JN
6749
25JN
6749
59JN
6749
41−
JN67
4999
EC
UA
DO
R
Ere
bida
eC
alpi
nae
Cal
yptr
a th
alic
tri
I,II
,III
,IV
,VM
M00
963
HQ
0061
56H
Q00
6861
HQ
0062
52H
Q00
6348
HQ
0067
63H
Q00
6435
HQ
0066
71H
Q00
6582
HQ
0069
55H
Q00
6504
TG
/TS
FIN
LA
ND
Ere
bida
eC
alpi
nae
Cal
yptr
a ho
kkai
daII
,III
RZ
336
JN40
1292
JN40
1174
JN40
1407
JN40
1518
JN40
0972
JN40
1616
JN40
1914
JN40
1823
JN40
1075
JN40
1718
TG
JAP
AN
Ere
bida
eC
alpi
nae
Cal
yptr
a la
taII
IR
Z43
1JN
6748
84JN
6748
66JN
6749
00JN
6749
17JN
6749
82JN
6749
31JN
6749
65JN
6749
47JN
6749
88−
TG
RU
SS
IA
Ere
bida
eC
alpi
nae
Cal
yptr
a ca
nade
nsis
III
CT
W2
−JN
6748
67−
−JN
6749
83JN
6749
33JN
6749
67JN
6749
49JN
6749
89JN
6750
05T
GU
SA
Ere
bida
eC
alpi
nae
Cal
yptr
a m
inut
icor
nis
III
RZ
514
JN67
4885
JN67
4868
JN67
4901
JN67
4918
JN67
4984
JN67
4932
JN67
4966
JN67
4948
JN67
4990
JN67
5006
TG
MA
LA
YS
IA
Ere
bida
eC
alpi
nae
Plu
siod
onta
nit
issi
ma
I,II
,III
RZ
333
HQ
0062
07H
Q00
6911
HQ
0063
03−
HQ
0068
15H
Q00
6470
HQ
0067
19H
Q00
6628
−−
CO
ST
A R
ICA
Ere
bida
eC
alpi
nae
Plu
siod
onta
coe
lono
taII
IR
Z10
6JN
6748
78JN
6748
60JN
6748
95JN
6749
11JN
6749
77JN
6749
26JN
6749
60JN
6749
42−
JN67
5000
HO
NG
KO
NG
Ere
bida
eC
alpi
nae
Plu
siod
onta
cas
taII
IR
Z42
9JN
6748
79JN
6748
61JN
6748
96JN
6749
12JN
6749
78JN
6749
27JN
6749
61JN
6749
43−
JN67
5001
RU
SS
IA
Ere
bida
eC
alpi
nae
Ora
esia
em
argi
nata
I,II
,III
RZ
102
HQ
0061
59H
Q00
6864
HQ
0062
56H
Q00
6351
HQ
0067
68H
Q00
6439
HQ
0066
75H
Q00
6586
HQ
0069
58H
Q00
6508
TS
HO
NG
KO
NG
Ere
bida
eC
alpi
nae
Ora
esia
exc
avat
aII
,III
RZ
337
JN40
1293
JN40
1175
JN40
1408
JN40
1519
JN40
0987
JN40
1617
JN40
1915
JN40
1824
JN40
1076
JN40
1719
HO
NG
KO
NG
Ere
bida
eC
alpi
nae
Ora
esia
exc
avat
eII
IR
Z43
4JN
6748
80JN
6748
62−
JN67
4913
−−
JN67
4958
−JN
6749
86−
US
A
Ere
bida
eC
alpi
nae
Ora
esia
nob
ilis
III
RZ
422
JN67
4881
JN67
4863
JN67
4897
JN67
4914
JN67
4979
JN67
4928
JN67
4962
JN67
4944
−JN
6750
02B
RA
ZIL
Ere
bida
eC
alpi
nae
Ora
esia
gla
ucoc
heil
aII
IR
Z41
8JN
6748
82JN
6748
64JN
6748
98JN
6749
15JN
6749
80JN
6749
29JN
6749
63JN
6749
45JN
6749
87JN
6750
03B
RA
ZIL
Ere
bida
eC
alpi
nae
Ora
esia
rec
tris
tria
III
RZ
433
JN67
4883
JN67
4865
JN67
4899
JN67
4916
JN67
4981
JN67
4930
JN67
4964
JN67
4946
−JN
6750
04N
EP
AL
Ere
bida
eH
ypoc
alin
aeH
ypso
roph
a ho
rmos
I,II
,III
RZ
17H
Q00
6176
HQ
0068
80H
Q00
6273
HQ
0063
67H
Q00
6784
HQ
0064
49H
Q00
6692
HQ
0066
00H
Q00
6972
HQ
0065
21U
SA
Ere
bida
eH
ypoc
alin
aeH
ypoc
ala
defl
orat
aII
RZ
105
JN40
1294
JN40
1176
JN40
1409
JN40
1520
JN40
0985
JN40
1618
JN40
1916
JN40
1825
JN40
1077
JN40
1720
TG
/TS
HO
NG
KO
NG
Ere
bida
eH
ypoc
alin
aeH
ypoc
ala
andr
emon
aII
,III
RZ
340
JN40
1295
JN40
1177
JN40
1410
JN40
1521
JN40
0980
JN40
1619
JN40
1917
JN40
1826
JN40
1078
JN40
1721
TG
CO
ST
A R
ICA
Ere
bida
eE
ulep
idot
inae
Tau
tobr
iga
glau
copi
sII
RZ
354
JN40
1296
JN40
1178
−−
JN40
0977
−−
JN40
1827
−−
CO
ST
A R
ICA
Ere
bida
eE
ulep
idot
inae
Pan
opod
a ru
fim
argo
I,II
,III
RZ
59H
Q00
6231
HQ
0069
34H
Q00
6326
HQ
0064
18H
Q00
6839
HQ
0064
88H
Q00
6743
HQ
0066
51H
Q00
7018
HQ
0065
64T
GU
SA
Ere
bida
eE
ulep
idot
inae
Ant
ible
mm
a fu
scir
etic
ulat
aII
,III
RZ
334
JN40
1297
JN40
1179
JN40
1411
−JN
4009
75JN
4016
20JN
4019
18JN
4018
28−
−C
OS
TA
RIC
A
Ere
bida
eE
ulep
idot
inae
Sany
s ir
rosc
aII
RZ
343
JN40
1298
JN40
1180
JN40
1412
−JN
4009
73JN
4016
21JN
4019
19JN
4018
29JN
4010
79JN
4017
22C
OS
TA
RIC
A
Ere
bida
eE
ulep
idot
inae
Eul
epid
otis
rec
tim
argo
I,II
,III
RZ
12H
Q00
6162
HQ
0069
60H
Q00
6259
HQ
0063
54H
Q00
6771
−H
Q00
6678
HQ
0065
88H
Q00
6960
HQ
0065
11T
GC
OS
TA
RIC
A
Ere
bida
eE
ulep
idot
inae
Ant
icar
sia
gem
mat
alis
IIR
Z26
7JN
4012
99JN
4011
81JN
4014
13JN
4015
22JN
4009
74JN
4016
22JN
4019
20JN
4018
30−
JN40
1723
TS
US
A
Ere
bida
eE
ulep
idot
inae
Ant
icar
sia
irro
rata
IIR
Z37
0JN
4013
00JN
4011
82JN
4014
14JN
4015
23JN
4009
95JN
4016
23JN
4019
21JN
4018
31JN
4010
80JN
4017
24H
ON
G K
ON
G
Ere
bida
eE
ulep
idot
inae
Hem
erop
lani
s fi
niti
ma
IIR
Z29
8JN
4013
01JN
4011
83JN
4014
15JN
4015
24JN
4010
04JN
4016
24JN
4019
22JN
4018
32−
JN40
1725
US
A
Ere
bida
eE
ulep
idot
inae
Oxi
derc
ia to
xea
I,II
RZ
295
HQ
0061
96H
Q00
6900
HQ
0062
92−
HQ
0068
04−
HQ
0067
09H
Q00
6617
HQ
0069
89−
TS
CO
ST
A R
ICA
Ere
bida
eE
ulep
idot
inae
Aze
ta c
eram
ina
I,II
,III
RZ
22H
Q00
6182
HQ
0068
86H
Q00
6278
HQ
0063
73H
Q00
6790
−H
Q00
6697
HQ
0066
05H
Q00
6978
HQ
0065
27C
OS
TA
RIC
A
Ere
bida
eT
oxoc
ampi
nae
Aut
ophi
la c
ham
aeph
anes
IIR
Z27
6JN
4013
02JN
4011
84JN
4014
16JN
4015
25JN
4010
05JN
4016
25JN
4019
23JN
4018
33JN
4010
81JN
4017
26C
OS
TA
RIC
A
Ere
bida
eT
oxoc
ampi
nae
Lyg
ephi
la p
asti
num
I,II
MM
0509
2G
U82
8711
GU
8285
06−
GU
8293
23G
U82
9587
−G
U83
0699
GU
8304
15G
U82
8199
GU
8300
97T
GFI
NL
AN
D
Ere
bida
eT
oxoc
ampi
nae
Lyg
ephi
la m
axim
aI,
IIR
Z57
HQ
0062
29H
Q00
6932
HQ
0063
24H
Q00
6416
HQ
0068
37H
Q00
6487
HQ
0067
41H
Q00
6649
−H
Q00
6562
TG
JAP
AN
Ere
bida
eT
inol
iinae
Tin
oliu
s eb
urne
igut
taII
RZ
331
JN40
1303
JN40
1185
JN40
1417
−−
−JN
4019
24JN
4018
34JN
4010
82−
TG
/TS
TH
AIL
AN
D
Ere
bida
eT
inol
iinae
Poe
ta d
enot
alis
IIR
Z44
5JN
4013
04JN
4011
86JN
4014
18JN
4015
26JN
4010
06JN
4016
26JN
4019
25−
−JN
4017
27T
SM
AL
AY
SIA
Ere
bida
eT
inol
iinae
Tam
sia
hier
ogly
phic
aII
RZ
389
JN40
1305
JN40
1187
JN40
1419
JN40
1527
JN40
1007
JN40
1627
JN40
1907
JN40
1835
JN40
1083
JN40
1728
MA
LA
YS
IA
Ere
bida
eU
nass
igne
dP
laty
jion
ia m
edio
rufa
IIR
Z11
1JN
4013
06JN
4011
88JN
4014
20JN
4015
28JN
4009
91JN
4016
28JN
4019
26−
JN40
1084
JN40
1729
HO
NG
KO
NG
Ere
bida
eSc
olec
ocam
pina
eSc
olec
ocam
pa li
burn
aI,
IIR
Z9
HQ
0062
42H
Q00
6944
HQ
0063
36H
Q00
6429
HQ
0068
50H
Q00
6495
HQ
0067
53H
Q00
6660
HQ
0070
25H
Q00
6573
TG
US
A
Appendix
41
Ere
bida
eSc
olec
ocam
pina
eG
abar
a st
ygia
lis
IIR
Z29
7JN
4013
07JN
4011
89JN
4014
21JN
4015
29−
JN40
1629
JN40
1927
JN40
1836
JN40
1085
JN40
1730
US
A
Ere
bida
eH
ypen
odin
aeH
ypen
odes
hum
idal
isI,
IIM
M01
780
GU
8286
71G
U82
8469
−G
U82
9285
GU
8295
56−
GU
8306
66−
GU
8281
68G
U83
0058
TG
/TS
FIN
LA
ND
Ere
bida
eH
ypen
odin
aeSc
hran
kia
cost
aest
riga
lis
I,II
RZ
27H
Q00
6192
HQ
0068
96H
Q00
6288
HQ
0063
82H
Q00
6800
HQ
0064
61H
Q00
6705
HQ
0066
13H
Q00
6987
−H
ON
G K
ON
G
Ere
bida
eH
ypen
odin
aeL
ucer
ia s
tria
taII
RZ
42JN
4013
08JN
4011
90JN
4014
22JN
4015
30JN
4010
08−
JN40
1928
−−
−H
ON
G K
ON
G
Ere
bida
eH
ypen
odin
aeL
ucer
ia o
cula
lis
IIR
Z36
9JN
4013
09JN
4011
91JN
4014
23JN
4015
31JN
4010
09−
JN40
1929
−JN
4010
86JN
4017
31H
ON
G K
ON
G
Ere
bida
eH
ypen
odin
aeA
nach
rost
is s
p.II
RZ
288
JN40
1310
JN40
1192
JN40
1424
−−
−JN
4019
30JN
4018
37JN
4010
87−
IND
ON
ES
IA
Ere
bida
eH
ypen
odin
aeM
icro
noct
ua s
p.I,
IIR
Z13
8H
Q00
6171
HQ
0068
75H
Q00
6268
HQ
0063
62H
Q00
6780
HQ
0064
45H
Q00
6687
HQ
0065
95H
Q00
6967
HQ
0065
16T
GIN
DO
NE
SIA
Ere
bida
eH
ypen
odin
aeB
iunc
us s
p. 1
IIR
Z47
5−
JN40
1193
JN40
1425
JN40
1532
JN40
0992
JN40
1630
JN40
1931
JN40
1838
JN40
1088
JN40
1732
GH
AN
A
Ere
bida
eH
ypen
odin
aeB
iunc
us s
p. 2
IIR
Z47
6JN
4013
11JN
4011
94JN
4014
26−
JN40
0994
JN40
1631
JN40
1908
JN40
1839
JN40
1089
JN40
1733
GH
AN
A
Ere
bida
eB
olet
obiin
aeSa
roba
pus
tuli
fera
I,II
RZ
104
HQ
0061
60H
Q00
6865
HQ
0062
57H
Q00
6352
HQ
0067
69−
HQ
0066
76−
−H
Q00
6509
TS
HO
NG
KO
NG
Ere
bida
eB
olet
obiin
aeC
onda
te s
p.II
RZ
393
JN40
1312
JN40
1195
JN40
1427
JN40
1533
JN40
1010
JN40
1632
JN40
1932
JN40
1840
JN40
1090
JN40
1734
MA
LA
YS
IA
Ere
bida
eB
olet
obiin
aeC
orga
tha
nite
nsI,
IIR
Z36
HQ
0062
11H
Q00
6915
HQ
0063
07H
Q00
6398
HQ
0068
19H
Q00
6474
HQ
0067
23H
Q00
6632
HQ
0070
01H
Q00
6547
HO
NG
KO
NG
Ere
bida
eB
olet
obiin
aeP
hyto
met
ra v
irid
aria
I,II
RZ
129
HQ
0061
65H
Q00
6962
HQ
0062
62H
Q00
6356
HQ
0067
74H
Q00
6442
HQ
0066
81H
Q00
6591
HQ
0069
62H
Q00
6512
TG
FIN
LA
ND
Ere
bida
eB
olet
obiin
aeLa
spey
ria
flexu
laI,
IIR
Z3
HQ
0061
97H
Q00
6901
HQ
0062
93H
Q00
6386
HQ
0068
05H
Q00
6463
HQ
0067
10H
Q00
6618
HQ
0069
90H
Q00
6536
TG
/TS
HU
NG
AR
Y
Ere
bida
eB
olet
obiin
aeZ
urob
ata
rora
taII
RZ
385
JN40
1313
JN40
1196
JN40
1428
JN40
1534
JN40
0996
−JN
4019
33JN
4018
41JN
4010
91JN
4017
35T
SM
AL
AY
SIA
Ere
bida
eB
olet
obiin
aeH
omod
es c
roce
aII
RZ
412
JN40
1314
JN40
1197
JN40
1429
JN40
1535
JN40
1011
−JN
4019
34JN
4018
42JN
4010
92JN
4017
36T
SM
AL
AY
SIA
Ere
bida
eB
olet
obiin
aeE
nisp
odes
pur
pure
aII
RZ
390
JN40
1315
JN40
1198
JN40
1430
−JN
4010
12JN
4016
33JN
4019
35JN
4018
43JN
4010
93JN
4017
37T
SM
AL
AY
SIA
Ere
bida
eB
olet
obiin
aeT
amba
mni
onom
era
IIR
Z41
5JN
4013
16JN
4011
99JN
4014
31JN
4015
36−
JN40
1634
JN40
1936
JN40
1844
−JN
4017
38M
AL
AY
SIA
Ere
bida
eB
olet
obiin
aeP
arol
ulis
abs
imil
isII
RZ
392
JN40
1317
JN40
1200
JN40
1432
JN40
1537
−JN
4016
35JN
4019
37JN
4018
45−
JN40
1739
MA
LA
YS
IA
Ere
bida
eB
olet
obiin
aeA
raeo
pter
on s
p.I,
IIR
Z13
7H
Q00
6170
HQ
0068
74H
Q00
6267
HQ
0063
61H
Q00
6779
−H
Q00
6686
−H
Q00
6966
HQ
0065
15T
GIN
DO
NE
SIA
Ere
bida
eB
olet
obiin
aeA
raeo
pter
on s
p.II
RZ
410
JN40
1318
JN40
1201
JN40
1433
JN40
1538
JN40
1013
JN40
1636
JN40
1938
JN40
1846
JN40
1094
−T
GM
AL
AY
SIA
Ere
bida
eB
olet
obiin
aeE
uble
mm
a pu
rpur
ina
I,II
RZ
7H
Q00
6237
HQ
0069
40H
Q00
6332
HQ
0064
24H
Q00
6845
HQ
0064
91H
Q00
6748
HQ
0066
55−
HQ
0065
69T
GH
UN
GA
RY
Ere
bida
eB
olet
obiin
aeE
uble
mm
a an
acho
resi
sII
RZ
98JN
4013
19JN
4012
02JN
4014
34JN
4015
39JN
4009
89JN
4016
37JN
4019
39JN
4018
47−
JN40
1741
TG
HO
NG
KO
NG
Ere
bida
eB
olet
obiin
aeE
uble
mm
a al
bifa
scia
IIR
Z22
0JN
4013
20JN
4012
03JN
4014
35JN
4015
40JN
4010
14JN
4016
38JN
4019
40−
−JN
4017
42T
GG
HA
NA
Ere
bida
eB
olet
obiin
aeP
aras
coti
afu
ligi
nari
aI,
IIM
M00
340
HQ
0061
54H
Q00
6862
HQ
0062
53H
Q00
6347
HQ
0067
64H
Q00
6436
HQ
0066
72H
Q00
6583
HQ
0069
54H
Q00
6505
TG
/TS
FIN
LA
ND
Ere
bida
eB
olet
obiin
aeM
etal
ectr
aed
ilis
IIR
Z37
2JN
4013
21JN
4012
04JN
4014
36JN
4015
41JN
4010
15JN
4016
39JN
4019
41JN
4018
48−
JN40
1744
FIN
LA
ND
Ere
bida
eB
olet
obiin
aeTr
isat
eles
em
ortu
alis
I,II
MM
0487
7G
U82
8707
GU
8285
02G
U82
9030
GU
8293
19G
U82
9583
GU
8298
21G
U83
0695
GU
8304
11G
U82
8195
GU
8300
93T
G/T
SFI
NL
AN
D
Ere
bida
eB
olet
obiin
aeP
rolo
phot
a tr
igon
ifer
aI,
IIR
Z37
HQ
0062
12H
Q00
6916
HQ
0063
08H
Q00
6399
HQ
0068
20H
Q00
6475
HQ
0067
24H
Q00
6633
HQ
0070
02−
TS
HO
NG
KO
NG
Ere
bida
eB
olet
obiin
aeH
ypen
agon
ia ?
brac
hypa
lpia
IIR
Z40
9JN
4013
22−
JN40
1437
JN40
1542
JN40
1016
JN40
1640
JN40
1942
JN40
1849
−−
MA
LA
YS
IA
Ere
bida
eB
olet
obiin
aeM
etae
men
e at
rigu
ttaI,
IIR
Z41
HQ
0062
18H
Q00
6922
HQ
0063
14H
Q00
6405
HQ
0068
26H
Q00
6481
HQ
0067
30H
Q00
6639
HQ
0070
07H
Q00
6552
HO
NG
KO
NG
Ere
bida
eB
olet
obiin
aeM
atae
omer
ase
mia
lba
IIR
Z10
7JN
4013
23JN
4012
05JN
4014
38−
−JN
4016
41−
JN40
1850
JN40
1095
JN40
1745
HO
NG
KO
NG
Ere
bida
eE
rebi
nae
Eua
onti
a se
mir
ufa
IIR
Z28
5JN
4013
24JN
4012
06JN
4014
39JN
4015
43JN
4010
17JN
4016
42JN
4019
43−
JN40
1096
JN40
1746
TS
US
A
Ere
bida
eE
rebi
nae
Aca
ntho
lipe
s ci
rcum
data
I,II
RZ
248
HQ
0061
89H
Q00
6893
HQ
0062
85H
Q00
6379
HQ
0067
97−
HQ
0067
02−
HQ
0069
84H
Q00
6531
TG
UA
E
Ere
bida
eE
rebi
nae
Aca
ntho
lipe
s re
gula
ris
I,II
RZ
135
HQ
0061
68H
Q00
6872
HQ
0062
65H
Q00
6359
HQ
0067
77−
HQ
0066
84−
−−
TG
/TS
RU
SS
IA
Ere
bida
eE
rebi
nae
Hyp
ospi
la b
olin
oide
sII
RZ
116
JN40
1325
JN40
1207
JN40
1440
JN40
1544
JN40
0997
JN40
1643
JN40
1944
JN40
1851
−JN
4017
43T
SH
ON
G K
ON
G
Ere
bida
eE
rebi
nae
Ugi
a in
susp
ecta
I,II
RZ
45H
Q00
6221
HQ
0069
25−
HQ
0064
08H
Q00
6829
HQ
0064
84H
Q00
6733
HQ
0066
42H
Q00
7010
HQ
0065
55H
ON
G K
ON
G
Ere
bida
eE
rebi
nae
Ugi
odes
cin
erea
IIR
Z32
6JN
4013
26JN
4012
08−
−−
JN40
1644
JN40
1945
−JN
4010
97−
TS
GH
AN
A
Appendix
42
Ere
bida
eE
rebi
nae
Sypn
oide
s fu
mos
aI,
IIR
Z31
3H
Q00
6201
HQ
0069
05H
Q00
6297
HQ
0063
90H
Q00
6809
HQ
0064
66H
Q00
6714
HQ
0066
22H
Q00
6994
HQ
0065
39JA
PA
N
Ere
bida
eE
rebi
nae
Dad
dala
luci
lla
IIR
Z32
0JN
4013
27JN
4012
09JN
4014
41JN
4015
45JN
4009
98JN
4016
45JN
4019
46JN
4018
52JN
4010
98JN
4017
47JA
PA
N
Ere
bida
eE
rebi
nae
Cat
ephi
a al
chym
ista
I,II
RZ
127
HQ
0061
64H
Q00
6961
HQ
0062
61H
Q00
6355
HQ
0067
73H
Q00
6441
HQ
0066
80H
Q00
6590
HQ
0069
61−
TG
/TS
GE
RM
AN
Y
Ere
bida
eE
rebi
nae
Het
eran
assa
sp.
IIR
Z35
0JN
4013
28JN
4012
10JN
4014
42JN
4015
46JN
4010
02JN
4016
46JN
4019
47JN
4018
60−
JN40
1748
US
A
Ere
bida
eE
rebi
nae
Zal
e be
thun
eiII
RZ
270
JN40
1329
JN40
1211
JN40
1443
JN40
1547
JN40
1018
JN40
1647
JN40
1948
JN40
1853
−JN
4017
49U
SA
Ere
bida
eE
rebi
nae
Thy
sani
a ze
nobi
aI,
IIR
Z53
HQ
0062
25H
Q00
6928
HQ
0063
20H
Q00
6412
HQ
0068
33H
Q00
6486
HQ
0067
37H
Q00
6645
HQ
0070
14H
Q00
6558
TG
CO
ST
A R
ICA
Ere
bida
eE
rebi
nae
Tox
onpr
ucha
sp.
IIR
Z30
7JN
4013
30JN
4012
12JN
4014
44JN
4015
48−
JN40
1648
JN40
1949
JN40
1854
−JN
4017
50U
SA
Ere
bida
eE
rebi
nae
Pse
udba
rydi
a cr
espu
laII
RZ
91JN
4013
31JN
4012
13JN
4014
45JN
4015
49JN
4010
19JN
4016
49JN
4019
50JN
4018
55JN
4010
99JN
4017
51C
OS
TA
RIC
A
Ere
bida
eE
rebi
nae
Pan
desm
a ro
bust
aI,
IIR
Z32
1H
Q00
6204
HQ
0069
08H
Q00
6300
HQ
0063
93H
Q00
6812
−H
Q00
6717
HQ
0066
25H
Q00
6997
HQ
0065
42T
G/T
SS
PA
IN
Ere
bida
eE
rebi
nae
Het
erop
alpi
a ac
rost
icta
I,II
RZ
243
HQ
0061
86H
Q00
6890
HQ
0062
82H
Q00
6376
HQ
0067
94−
HQ
0067
00−
HQ
0069
81−
UA
E
Ere
bida
eE
rebi
nae
Sphi
ngom
orph
a ch
lore
aI,
IIR
Z29
1H
Q00
6195
HQ
0068
99H
Q00
6291
HQ
0063
85H
Q00
6803
−H
Q00
6708
HQ
0066
16−
−T
ST
AN
ZA
NIA
Ere
bida
eE
rebi
nae
Per
icym
a cr
uege
riI,
IIR
Z99
HQ
0062
44H
Q00
6946
HQ
0063
38H
Q00
6431
HQ
0068
52H
Q00
6497
HQ
0067
55H
Q00
6662
HQ
0070
27H
Q00
6575
TG
HO
NG
KO
NG
Ere
bida
eE
rebi
nae
Sym
pis
rufi
basi
sI,
IIR
Z48
HQ
0062
23−
HQ
0063
18H
Q00
6410
HQ
0068
31H
Q00
6485
HQ
0067
35H
Q00
6644
HQ
0070
12H
Q00
6557
TS
HO
NG
KO
NG
Ere
bida
eE
rebi
nae
Ere
bus
ephe
sper
isI,
IIR
Z11
HQ
0061
61H
Q00
6866
HQ
0062
58H
Q00
6353
HQ
0067
70H
Q00
6440
HQ
0066
77H
Q00
6587
HQ
0069
59H
Q00
6510
TG
TA
IWA
N
Ere
bida
eE
rebi
nae
Ery
gia
apic
alis
I,II
RZ
29H
Q00
6194
HQ
0068
98H
Q00
6290
HQ
0063
84H
Q00
6802
−H
Q00
6707
HQ
0066
15H
Q00
6988
HQ
0065
35T
SH
ON
G K
ON
G
Ere
bida
eE
rebi
nae
Bul
ia d
educ
taII
RZ
314
JN40
1332
JN40
1214
JN40
1446
JN40
1550
JN40
1020
JN40
1650
JN40
1951
JN40
1856
JN40
1100
JN40
1752
US
A
Ere
bida
eE
rebi
nae
For
sebi
a pe
rlae
taII
RZ
284
JN40
1333
JN40
1215
JN40
1447
JN40
1551
JN40
1021
JN40
1651
JN40
1952
JN40
1857
JN40
1101
JN40
1753
TS
US
A
Ere
bida
eE
rebi
nae
Mel
ipot
is p
unct
ifini
sII
RZ
342
JN40
1334
JN40
1216
JN40
1448
JN40
1552
JN40
1022
JN40
1652
JN40
1953
JN40
1858
JN40
1102
JN40
1754
TG
CO
ST
A R
ICA
Ere
bida
eE
rebi
nae
Mel
ipot
is ju
cund
aI,
IIR
Z58
HQ
0062
30H
Q00
6933
HQ
0063
25H
Q00
6417
HQ
0068
38−
HQ
0067
42H
Q00
6650
HQ
0070
17H
Q00
6563
TG
/TS
US
A
Ere
bida
eE
rebi
nae
Pho
beri
a at
omar
isII
RZ
286
JN40
1335
JN40
1217
JN40
1449
−JN
4010
23−
JN40
1954
JN40
1859
JN40
1103
JN40
1755
TS
US
A
Ere
bida
eE
rebi
nae
Aud
ea b
ipun
ctat
aI,
IIR
Z60
HQ
0062
33H
Q00
6936
HQ
0063
28H
Q00
6420
HQ
0068
41−
HQ
0067
44H
Q00
6652
HQ
0070
19H
Q00
6565
TG
/TS
CO
NG
O
Ere
bida
eE
rebi
nae
Aud
ea h
umer
alis
IIR
Z29
0JN
4013
36JN
4012
18JN
4014
50JN
4015
53JN
4010
24JN
4016
53JN
4019
55−
JN40
1104
JN40
1756
TG
TA
NZ
AN
IA
Ere
bida
eE
rebi
nae
Hyp
otac
ha b
rand
berg
ensi
sII
RZ
275
JN40
1337
JN40
1219
JN40
1451
JN40
1554
JN40
0999
JN40
1654
JN40
1956
−JN
4011
05JN
4017
57N
AM
IBIA
Ere
bida
eE
rebi
nae
Cat
ocal
a sp
onsa
I,II
,IV
,VM
M04
358
GU
8287
00G
U82
8495
GU
8290
23G
U82
9312
GU
8295
76G
U82
9816
GU
8306
88G
U83
0404
GU
8281
89G
U83
0086
TG
FIN
LA
ND
Ere
bida
eE
rebi
nae
Ulo
tric
hopu
s m
acul
aI,
IIR
Z24
1H
Q00
6185
HQ
0068
89H
Q00
6281
−H
Q00
6793
HQ
0064
57H
Q00
6699
HQ
0066
08−
HQ
0065
30T
AIW
AN
Ere
bida
eE
rebi
nae
Hyp
opyr
a ca
pens
isII
RZ
149
HQ
0061
72H
Q00
6876
HQ
0062
69H
Q00
6363
HQ
0067
81−
HQ
0066
88H
Q00
6596
HQ
0069
68H
Q00
6517
TG
GH
AN
A
Ere
bida
eE
rebi
nae
Spir
ama
reto
rta
IIR
Z35
9JN
4013
38JN
4012
20JN
4014
52−
JN40
1025
JN40
1655
JN40
1957
−JN
4011
06JN
4017
58T
AIW
AN
Ere
bida
eE
rebi
nae
Cal
ypti
s id
onea
IIR
Z47
3JN
4013
39JN
4012
21JN
4014
53−
JN40
1000
JN40
1656
JN40
1958
JN40
1861
JN40
1107
JN40
1759
EC
UA
DO
R
Ere
bida
eE
rebi
nae
Om
mat
opho
ra lu
min
osa
IIR
Z40
7JN
4013
40JN
4012
22JN
4014
54JN
4015
55JN
4010
26JN
4016
57JN
4019
59JN
4018
62JN
4011
08JN
4017
60T
G/T
SM
AL
AY
SIA
Ere
bida
eE
rebi
nae
Pan
tydi
adi
emen
iI,
IIR
Z30
9H
Q00
6199
HQ
0069
03H
Q00
6295
HQ
0063
88H
Q00
6807
HQ
0064
64H
Q00
6712
HQ
0066
20H
Q00
6992
HQ
0065
38A
US
TR
AL
IA
Ere
bida
eE
rebi
nae
Moc
is la
tipes
I,II
RZ
20H
Q00
6180
HQ
0068
84H
Q00
6276
HQ
0063
71H
Q00
6788
HQ
0064
53H
Q00
6695
HQ
0066
03H
Q00
6976
HQ
0065
25C
OS
TA
RIC
A
Ere
bida
eE
rebi
nae
Cal
list
ege
mi
I,II
MM
0546
9H
Q00
6150
HQ
0068
57H
Q00
6248
HQ
0063
43H
Q00
6759
−H
Q00
6667
HQ
0065
78H
Q00
6950
HQ
0065
00T
SFI
NL
AN
D
Ere
bida
eE
rebi
nae
Euc
lidi
a gl
yphi
caI,
IIR
Z82
HQ
0062
39H
Q00
6942
HQ
0063
33H
Q00
6426
HQ
0068
47−
HQ
0067
50H
Q00
6657
HQ
0070
23H
Q00
6570
TG
FIN
LA
ND
Ere
bida
eE
rebi
nae
Erc
heia
cyl
lari
aI,
IIR
Z33
HQ
0062
05H
Q00
6909
HQ
0063
01H
Q00
6394
HQ
0068
13−
HQ
0067
18H
Q00
6626
HQ
0069
98H
Q00
6543
TG
HO
NG
KO
NG
Ere
bida
eE
rebi
nae
Hul
odes
car
anea
I,II
RZ
126
HQ
0061
63−
HQ
0062
60−
HQ
0067
72−
HQ
0066
79H
Q00
6589
−−
TG
MA
LA
YS
IA
Ere
bida
eE
rebi
nae
Eri
ceia
sub
cine
rea
IIR
Z39
HQ
0062
14H
Q00
6918
HQ
0063
10H
Q00
6401
HQ
0068
22H
Q00
6477
HQ
0067
26H
Q00
6635
HQ
0070
04H
Q00
6549
HO
NG
KO
NG
Ere
bida
eE
rebi
nae
Pla
tyja
um
min
eaII
RZ
261
JN40
1341
JN40
1223
JN40
1455
JN40
1556
JN40
1027
JN40
1658
JN40
1960
JN40
1863
−JN
4017
61T
SH
ON
G K
ON
G
Ere
bida
eE
rebi
nae
Ani
sone
ura
sale
bros
aI,
IIR
Z38
HQ
0062
13H
Q00
6917
HQ
0063
09H
Q00
6400
HQ
0068
21H
Q00
6476
HQ
0067
25−
HQ
0070
03H
Q00
6548
TS
HO
NG
KO
NG
Appendix
43
Ere
bida
eE
rebi
nae
Pra
xis
porp
hyre
tica
IIR
Z30
8JN
4013
42JN
4012
24JN
4014
56−
JN40
1028
JN40
1659
JN40
1961
JN40
1864
JN40
1109
JN40
1762
AU
ST
RA
LIA
Ere
bida
eE
rebi
nae
Isch
yja
man
liaII
RZ
269
JN40
1343
JN40
1225
JN40
1457
JN40
1557
JN40
1029
JN40
1660
JN40
1962
JN40
1822
JN40
1110
JN40
1763
TS
MA
LA
YS
IA
Ere
bida
eE
rebi
nae
Oxy
odes
scr
obic
ulat
aII
RZ
113
JN40
1344
JN40
1226
JN40
1458
JN40
1558
JN40
1030
JN40
1661
JN40
1963
JN40
1866
JN40
1111
JN40
1764
TS
HO
NG
KO
NG
Ere
bida
eE
rebi
nae
Serr
odes
cam
pana
I,II
RZ
318
HQ
0062
02H
Q00
6906
HQ
0062
98H
Q00
6391
HQ
0068
10H
Q00
6467
HQ
0067
15H
Q00
6623
HQ
0069
95H
Q00
6540
TA
IWA
N
Ere
bida
eE
rebi
nae
Ava
tha
ulop
tera
IIR
Z31
7JN
4013
45JN
4012
27JN
4014
59JN
4015
59JN
4010
03JN
4016
62JN
4019
64JN
4018
67−
JN40
1765
MA
LA
YS
IA
Ere
bida
eE
rebi
nae
Coc
ytia
dur
vill
iiII
RZ
401
JN40
1346
JN40
1228
JN40
1460
JN40
1560
JN40
1031
JN40
1663
JN40
1965
JN40
1869
JN40
1112
JN40
1766
TG
/TS
NE
W G
UIN
EA
Ere
bida
eE
rebi
nae
Bas
till
a pr
aete
rmis
saII
RZ
306
JN40
1347
JN40
1229
JN40
1461
JN40
1561
JN40
1032
JN40
1664
JN40
1966
JN40
1868
JN40
1113
JN40
1767
MA
LA
YS
IA
Ere
bida
eE
rebi
nae
Ach
aea
serv
aI,
IIR
Z19
HQ
0061
79H
Q00
6883
HQ
0062
75H
Q00
6370
HQ
0067
87H
Q00
6452
HQ
0066
94H
Q00
6602
HQ
0069
75H
Q00
6524
MA
LA
YS
IA
Ere
bida
eE
rebi
nae
Cha
lcio
pe m
ygdo
nII
RZ
391
JN40
1348
JN40
1230
JN40
1462
JN40
1562
JN40
1001
JN40
1665
JN40
1967
JN40
1865
JN40
1881
JN40
1768
TS
MA
LA
YS
IA
Ere
bida
eE
rebi
nae
All
otri
a el
onym
pha
IIR
Z29
4JN
4013
49JN
4012
31JN
4014
63JN
4015
63JN
4010
33JN
4016
66JN
4019
68JN
4018
70JN
4011
15JN
4017
69T
SU
SA
Ere
bida
eE
rebi
nae
Cly
tie d
evia
I,II
RZ
247
HQ
0061
88H
Q00
6892
HQ
0062
84H
Q00
6378
HQ
0067
96H
Q00
6459
−H
Q00
6610
HQ
0069
83−
UA
E
Ere
bida
eE
rebi
nae
Oph
iusa
tirh
aca
I,II
,IV
RZ
246
HQ
0061
87H
Q00
6891
HQ
0062
83H
Q00
6377
HQ
0067
95H
Q00
6458
HQ
0067
01H
Q00
6609
HQ
0069
82−
TG
/TS
UA
E
Ere
bida
eE
rebi
nae
Thy
as m
etap
haea
IIR
Z19
0JN
4013
50JN
4012
32JN
4014
64JN
4015
64JN
4010
34JN
4016
67JN
4019
69JN
4018
71JN
4011
16JN
4017
70G
HA
NA
Ere
bida
eE
rebi
nae
Oph
iusa
cor
onat
aI,
IIR
Z21
HQ
0061
81H
Q00
6885
HQ
0062
77H
Q00
6372
HQ
0067
89H
Q00
6454
HQ
0066
96H
Q00
6604
HQ
0069
77H
Q00
6526
TG
MA
LA
YS
IA
Ere
bida
eE
rebi
nae
Art
ena
dota
taI,
IIR
Z46
HQ
0062
22H
Q00
6926
HQ
0063
17H
Q00
6409
HQ
0068
30−
HQ
0067
34H
Q00
6643
HQ
0070
11H
Q00
6556
HO
NG
KO
NG
Nol
idae
Dip
hthe
rina
eL
epid
odes
gal
lopa
voIV
RZ
353
XX
XX
X−
−X
X−
CO
ST
A R
ICA
Nol
idae
Dip
hthe
rina
eL
epid
odes
lim
bula
ta 1
IVR
Z62
8−
−X
XX
X−
XX
XT
SC
OS
TA
RIC
A
Nol
idae
Dip
hthe
rina
eL
epid
odes
lim
bula
ta 2
IVR
Z63
0X
−X
XX
X−
XX
XT
SC
OS
TA
RIC
A
Nol
idae
Dip
hthe
rina
eL
epid
odes
lim
bula
ta 3
IVR
Z59
4X
X−
−X
−−
−−
−T
SC
OS
TA
RIC
A
Nol
idae
Dip
hthe
rina
eD
ipht
hera
fest
iva
2IV
RZ
465
XX
XX
XX
−X
XX
TG
/TS
CO
ST
A R
ICA
Nol
idae
Dip
hthe
rina
eD
ipht
hera
fest
iva
3IV
RZ
631
X−
−−
X−
X−
−−
TG
/TS
CO
ST
A R
ICA
Nol
idae
Dip
hthe
rina
eD
ipht
hera
fest
iva
1IV
RZ
633
X−
X−
X−
−−
−−
TG
/TS
CO
ST
A R
ICA
Nol
idae
Ris
obin
aeR
isob
a ob
stru
cta
2II
,IV
RZ
381
JN40
1233
JN40
1117
JN40
1351
−JN
4009
26JN
4015
65JN
4018
72JN
4017
71JN
4010
35JN
4016
68T
GM
AL
AY
SIA
Nol
idae
Ris
obin
aeR
isob
a ob
stru
cta
1I,
II,I
VR
Z65
XX
XX
XX
XX
−X
TG
TH
AIL
AN
D
Nol
idae
Ris
obin
aeB
aile
ya le
vita
nsIV
RZ
461
XX
XX
XX
XX
XX
US
A
Nol
idae
Col
lom
enin
aeC
ollo
men
a si
oper
a 1
IVR
Z63
2X
−−
−X
−X
X−
−T
GC
OS
TA
RIC
A
Nol
idae
Col
lom
enin
aeC
ollo
men
a si
oper
a 2
IV,V
RZ
634
X−
XX
XX
XX
−−
TG
CO
ST
A R
ICA
Nol
idae
Col
lom
enin
aeN
eost
icto
pera
nig
ropu
ncta
IV,V
RZ
469
XX
XX
XX
XX
XX
TS
CO
ST
A R
ICA
Nol
idae
Bea
nina
eB
eana
term
inig
era
IVR
Z63
8X
−X
XX
XX
XX
XT
GT
HA
ILA
ND
Nol
idae
Elig
min
aeB
aroa
sia
mic
aIV
RZ
396
XX
XX
XX
XX
XX
MA
LA
YS
IA
Nol
idae
Elig
min
aeSe
lepa
mol
ybde
aI,
IVR
Z32
HQ
0062
03H
Q00
6907
HQ
0062
99H
Q00
6392
HQ
0068
11H
Q00
6468
HQ
0067
16H
Q00
6624
HQ
0069
96H
Q00
6541
HO
NG
KO
NG
Nol
idae
Elig
min
aeSe
lepa
dis
cige
ra/c
elti
sIV
RZ
500
XX
XX
X−
XX
XX
HO
NG
KO
NG
Nol
idae
Elig
min
aeP
tisc
iana
sem
iniv
eaIV
RZ
516
XX
XX
XX
XX
−X
TS
MA
LA
YS
IA
Nol
idae
Elig
min
aeT
rior
bis
aure
ovitt
aIV
RZ
360
XX
XX
XX
XX
XX
MA
LA
YS
IA
Nol
idae
Elig
min
aeE
ligm
a na
rcis
sus
I,II
,IV
,VR
Z97
HQ
0062
43H
Q00
6945
HQ
0063
37H
Q00
6430
HQ
0068
51H
Q00
6496
HQ
0067
54H
Q00
6661
HQ
0070
26H
Q00
6574
TG
/TS
HO
NG
KO
NG
Nol
idae
Elig
min
aeE
laeo
gnat
ha a
rgyr
itis
IVR
Z46
2X
XX
−X
XX
X−
−T
SC
OS
TA
RIC
A
Nol
idae
Elig
min
aeIs
cadi
a pr
oduc
tIV
RZ
361
XX
−−
X−
XX
−−
CO
ST
A R
ICA
Nol
idae
Wes
term
anni
inae
Pte
rogo
nia
card
inal
isIV
RZ
522
XX
XX
XX
XX
X−
MA
LA
YS
IA
Appendix
44
Nol
idae
Wes
term
anni
inae
Mia
rom
ima
corn
ucop
iaIV
RZ
530
XX
XX
XX
XX
−X
MA
LA
YS
IA
Nol
idae
Wes
term
anni
inae
Wes
term
anni
a su
perb
aIV
RZ
645
X−
XX
XX
XX
X−
TG
/TS
TH
AIL
AN
D
Nol
idae
Wes
term
anni
inae
Irag
aode
s no
bili
sIV
RZ
483
XX
XX
XX
X−
−−
TS
JAP
AN
Nol
idae
Wes
term
anni
inae
Neg
eta
cont
rari
ata
2I,
II,I
VR
Z26
HQ
0061
91H
Q00
6895
HQ
0062
87H
Q00
6381
HQ
0067
99H
Q00
6460
HQ
0067
04H
Q00
6612
HQ
0069
86H
Q00
6533
HO
NG
KO
NG
Nol
idae
Wes
term
anni
inae
Neg
eta
abbr
evia
taIV
RZ
414
X−
XX
XX
XX
XX
MA
LA
YS
IA
Nol
idae
Wes
term
anni
inae
Neg
eta
cont
rari
ata
1IV
RZ
487
XX
X−
XX
XX
−X
TA
IWA
N
Nol
idae
Wes
term
anni
inae
Neg
eta
cont
rari
ata
3IV
RZ
504
XX
XX
XX
XX
XX
TS
MA
LA
YS
IA
Nol
idae
Nol
inae
Mel
anog
raph
ia fl
exil
inea
taIV
RZ
498
XX
XX
XX
XX
XX
TS
HO
NG
KO
NG
Nol
idae
Nol
inae
Mem
bran
ola
lam
pang
IVR
Z62
6X
XX
XX
XX
X−
XT
HA
ILA
ND
Nol
idae
Nol
inae
Alc
anol
a ty
mpa
nist
esIV
RZ
624
XX
XX
XX
XX
XX
TH
AIL
AN
D
Nol
idae
Nol
inae
Alc
anol
a ob
scur
ata
IVR
Z62
5X
−X
XX
XX
XX
−T
HA
ILA
ND
Nol
idae
Nol
inae
Meg
anol
a m
inus
cula
IVR
Z48
5X
XX
XX
XX
−−
XU
SA
Nol
idae
Nol
inae
Nol
a ba
sini
gra
IVR
Z62
7X
XX
XX
XX
XX
XT
GT
HA
ILA
ND
Nol
idae
Nol
inae
unkn
own
nolin
eIV
RZ
526
XX
XX
XX
X−
XX
MA
LA
YS
IA
Nol
idae
Nol
inae
Nol
a ae
rugu
laI,
II,I
VM
M01
776
GU
8286
70G
U82
8468
GU
8290
01G
U82
9284
GU
8295
55−
GU
8306
65G
U83
0376
GU
8281
67G
U83
0057
TG
FIN
LA
ND
Nol
idae
Nol
inae
Nol
a m
inna
IVR
Z48
8X
XX
XX
XX
X−
XT
GU
SA
Nol
idae
Nol
inae
Nol
a cr
etac
eaIV
RZ
493
XX
X−
XX
X−
X−
TG
HO
NG
KO
NG
Nol
idae
Nol
inae
Nol
a an
alis
IVR
Z49
4X
X−
XX
X−
−X
−T
GH
ON
G K
ON
G
Nol
idae
Nol
inae
Nol
a lu
cida
lis
IVR
Z53
3X
XX
XX
−X
X−
−T
GM
AL
AY
SIA
Nol
idae
Nol
inae
Nol
a fa
scia
taIV
RZ
534
X−
XX
XX
XX
−X
TG
MA
LA
YS
IA
Nol
idae
Nol
inae
Man
oba
brun
ellu
sIV
RZ
495
XX
XX
XX
−X
XX
HO
NG
KO
NG
Nol
idae
Chl
oeph
orin
aeM
aced
a m
ansu
eta
IVR
Z64
1X
−−
−X
−X
X−
−T
ST
HA
ILA
ND
Nol
idae
Chl
oeph
orin
aeP
arac
ram
a an
gula
taI,
IVR
Z43
HQ
0062
19H
Q00
6923
HQ
0063
15H
Q00
6406
HQ
0068
27H
Q00
6482
HQ
0067
31H
Q00
6640
HQ
0070
08H
Q00
6553
HO
NG
KO
NG
Nol
idae
Chl
oeph
orin
aeP
arac
ram
a du
lcis
sim
aIV
RZ
648
X−
XX
XX
XX
−−
TS
TH
AIL
AN
D
Nol
idae
Chl
oeph
orin
aeE
aria
s cl
oran
aI,
II,I
VM
M06
650
GU
8287
47G
U92
9721
GU
8290
61G
U82
9346
GU
8296
10G
U82
9845
GU
8307
18G
U83
0448
GU
8282
31G
U83
0131
TG
/TS
FIN
LA
ND
Nol
idae
Chl
oeph
orin
aeE
aria
s ro
seife
raIV
RZ
480
XX
XX
XX
XX
XX
TA
IWA
N
Nol
idae
Chl
oeph
orin
aeE
aria
s in
sula
naIV
RZ
492
XX
X−
XX
−X
XX
TG
HO
NG
KO
NG
Nol
idae
Chl
oeph
orin
aeM
auri
lia
icon
ica
IVR
Z48
4X
X−
XX
XX
XX
XIN
DO
NE
SIA
Nol
idae
Chl
oeph
orin
aeTo
rtri
cifo
rma
razo
wsk
iiIV
RZ
586
XX
XX
XX
XX
−−
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeD
idig
ua v
irid
ifas
cia
IVR
Z52
5X
XX
XX
XX
X−
XM
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeX
enoc
hroa
ann
aeIV
RZ
386
XX
XX
XX
XX
XX
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeX
enoc
hroa
xan
thia
IVR
Z50
3X
XX
XX
XX
XX
XM
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeD
iehl
ea tu
mid
aIV
RZ
479
XX
XX
X−
XX
X−
TS
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeC
hlor
iola
gra
tiss
ima
IVR
Z53
1X
XX
XX
XX
XX
XT
SM
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeG
elas
toce
ra v
irid
imac
ula
IVR
Z52
1X
XX
XX
XX
X−
XT
GM
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeB
eara
tort
rici
form
isIV
RZ
515
XX
XX
XX
XX
−X
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeA
riol
ica
arge
ntea
I,IV
RZ
63H
Q00
6234
HQ
0069
37H
Q00
6329
HQ
0064
21H
Q00
6842
−H
Q00
6745
HQ
0066
53H
Q00
7020
HQ
0065
66T
GJA
PA
N
Nol
idae
Chl
oeph
orin
aeG
arig
a m
irab
ilis
IVR
Z11
4X
XX
−X
XX
X−
−H
ON
G K
ON
G
Appendix
45
Nol
idae
Chl
oeph
orin
aeT
ympa
nist
es fu
sim
argo
IVR
Z56
6X
XX
XX
XX
XX
−M
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeP
seud
oips
pra
sina
na 2
IVR
Z12
8X
−X
XX
XX
−−
−T
G/T
SG
ER
MA
NY
Nol
idae
Chl
oeph
orin
aeP
seud
oips
pra
sina
na 1
I,IV
,VM
M00
107
GU
8286
00G
U82
8399
GU
8289
34G
U82
9229
GU
8294
96G
U82
9754
GU
8306
11G
U83
0312
GU
8281
01G
U82
9989
TG
FIN
LA
ND
Nol
idae
Chl
oeph
orin
aeC
leth
roph
ora
dist
inct
aIV
RZ
478
XX
XX
XX
XX
XX
TS
JAP
AN
Nol
idae
Chl
oeph
orin
aeH
ylop
hilo
des
orie
ntal
isIV
RZ
643
X−
XX
X−
XX
X−
TS
TH
AIL
AN
D
Nol
idae
Chl
oeph
orin
aeC
hlor
opla
ga s
p.IV
RZ
520
XX
XX
XX
XX
XX
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeC
hlor
opla
ga n
ygm
iaIV
RZ
646
X−
XX
XX
XX
−X
TS
TH
AIL
AN
D
Nol
idae
Chl
oeph
orin
aeSi
nna
extr
ema
IVR
Z49
0X
XX
XX
XX
XX
XJA
PA
N
Nol
idae
Chl
oeph
orin
aeSi
nna
flor
alis
IVR
Z52
9X
XX
XX
XX
X−
XM
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeSi
glop
hora
ferr
eilu
tea
IVR
Z48
9X
XX
XX
XX
XX
−JA
PA
N
Nol
idae
Chl
oeph
orin
aeC
hand
ica
quad
ripe
nnis
IVR
Z57
9X
XX
XX
XX
X−
XT
SM
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeC
osse
dia
hyri
odes
IVR
Z52
4X
XX
XX
XX
X−
−M
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeLa
band
a ce
ylus
alis
IVR
Z55
5X
XX
XX
XX
X−
XM
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeA
quis
orb
icul
aris
IVR
Z63
9X
−X
XX
XX
X−
−T
HA
ILA
ND
Nol
idae
Chl
oeph
orin
aeT
atho
thri
pa c
onti
nua
IVR
Z64
0−
−X
XX
XX
X−
−T
ST
HA
ILA
ND
Nol
idae
Chl
oeph
orin
aeB
leni
na d
onan
sI,
IVR
Z64
HQ
0062
35H
Q00
6938
HQ
0063
30H
Q00
6422
HQ
0068
43H
Q00
6490
HQ
0067
46H
Q00
6654
HQ
0070
21H
Q00
6567
TG
SU
MA
TR
A
Nol
idae
Chl
oeph
orin
aeB
leni
na q
uina
ria
IVR
Z49
9X
XX
XX
XX
XX
XT
GH
ON
G K
ON
G
Nol
idae
Chl
oeph
orin
aeD
ilop
hoth
ripa
chr
ysor
rhae
aIV
RZ
528
XX
XX
XX
XX
−−
TS
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeE
gchi
rete
s no
mim
usIV
RZ
481
XX
XX
X−
X−
−−
TS
CO
ST
A R
ICA
Nol
idae
Chl
oeph
orin
aeN
ycte
ola
dege
nera
naI,
IVM
M00
135
GU
8286
12G
U82
8410
GU
8289
46G
U82
9238
GU
8295
06G
U82
9760
GU
8306
21G
U83
0323
GU
8281
13G
U83
0000
TG
FIN
LA
ND
Nol
idae
Chl
oeph
orin
aeN
ycte
ola
sinu
osa
IVR
Z50
1X
XX
XX
XX
X−
−T
GH
ON
G K
ON
G
Nol
idae
Chl
oeph
orin
aeN
ycte
ola
indi
caIV
RZ
539
XX
XX
XX
XX
−X
TG
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeG
arel
la n
ilot
ica
IVR
Z47
7X
XX
XX
XX
XX
XU
SA
Nol
idae
Chl
oeph
orin
aeG
arel
la r
ufic
irra
IVR
Z50
2X
XX
XX
−X
X−
XH
ON
G K
ON
G
Nol
idae
Chl
oeph
orin
aeG
iaur
a ro
bust
aI,
IVR
Z31
HQ
0062
00H
Q00
6904
HQ
0062
96H
Q00
6389
HQ
0068
08H
Q00
6465
HQ
0067
13H
Q00
6621
HQ
0069
93−
HO
NG
KO
NG
Nol
idae
Chl
oeph
orin
aeG
iaur
a ni
gril
inea
taIV
RZ
519
X−
XX
XX
XX
−X
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeG
iaur
a m
ulti
punc
tata
IVR
Z49
6X
XX
XX
XX
XX
XH
ON
G K
ON
G
Nol
idae
Chl
oeph
orin
aeG
iaur
a ni
veid
isca
IVR
Z52
7X
XX
XX
XX
X−
XM
AL
AY
SIA
Nol
idae
Chl
oeph
orin
aeP
arda
sena
ver
naIV
RZ
532
XX
XX
X−
XX
X−
MA
LA
YS
IA
Nol
idae
Chl
oeph
orin
aeE
tann
a cl
opae
aIV
RZ
486
XX
XX
XX
XX
−X
AU
ST
RA
LIA
Nol
idae
Chl
oeph
orin
aeE
tann
a br
eviu
scul
a 2
IVR
Z49
1X
XX
XX
XX
XX
XH
ON
G K
ON
G
Nol
idae
Chl
oeph
orin
aeE
tann
a br
eviu
scul
a 1
IVR
Z49
7X
XX
XX
XX
X−
XH
ON
G K
ON
G
Nol
idae
Chl
oeph
orin
aeE
tann
a br
eviu
scul
a 3
IVR
Z51
8X
XX
XX
XX
X−
XM
AL
AY
SIA
Noc
tuid
aeD
yops
inae
Dyo
ps c
hrom
atop
hila
I,II
,VR
Z10
HQ
0061
58−
HQ
0062
55H
Q00
6350
HQ
0067
67H
Q00
6438
HQ
0066
74H
Q00
6585
HQ
0069
57H
Q00
6507
TG
CO
ST
A R
ICA
Noc
tuid
aeD
yops
inae
Pse
udoa
rcte
mel
anis
II,V
RZ
147
JN40
1235
JN40
1119
JN40
1353
JN40
1465
JN40
0928
JN40
1567
JN40
1874
JN40
1773
JN40
1037
JN40
1669
GH
AN
A
Noc
tuid
aeD
yops
inae
Arc
te m
odes
taI,
II,V
RZ
54H
Q00
6226
HQ
0069
29H
Q00
6321
HQ
0064
13H
Q00
6834
−H
Q00
6738
HQ
0066
46H
Q00
7015
HQ
0065
59T
GM
AL
AY
SIA
Noc
tuid
aeD
yops
inae
Par
arct
e sc
hnei
deri
ana
VR
Z54
8X
XX
XX
XX
X−
−T
SC
OS
TA
RIC
A
Noc
tuid
aeD
yops
inae
Bel
cian
a ka
laII
,VR
Z41
6JN
4012
38−
JN40
1356
JN40
1468
JN40
0930
JN40
1570
JN40
1879
JN40
1776
JN40
1040
JN40
1672
MA
LA
YS
IA
Appendix
46
Noc
tuid
aeD
yops
inae
Bel
cian
a bi
form
isII
,VR
Z38
4JN
4012
37JN
4011
21JN
4013
55JN
4014
67JN
4009
29JN
4015
69JN
4018
77JN
4017
75JN
4010
39JN
4016
71T
SM
AL
AY
SIA
Noc
tuid
aeD
yops
inae
Cyc
lode
s om
ma
VR
Z63
6X
XX
XX
XX
X−
XT
ST
HA
ILA
ND
Noc
tuid
aeD
yops
inae
Cer
octe
na a
myn
taV
RZ
552
XX
XX
XX
XX
X−
TS
EC
UA
DO
R
Noc
tuid
aeD
yops
inae
Sosx
etra
gra
taII
,VR
Z28
1JN
4012
36JN
4011
20JN
4013
54JN
4014
66JN
4009
88JN
4015
68JN
4018
75JN
4017
74JN
4010
38JN
4016
70T
SC
OS
TA
RIC
A
Noc
tuid
aePl
usiin
aeA
utog
raph
a ga
mm
aI,
II,I
V,V
MM
0032
8G
U82
8636
GU
8284
34G
U82
8970
GU
8292
56G
U82
9528
−G
U83
0640
GU
8303
44G
U82
8135
GU
8300
23T
G/T
SFI
NL
AN
D
Noc
tuid
aePl
usiin
aeA
bros
tola
trip
arti
taI,
VM
M05
132
HQ
0061
52H
Q00
6859
HQ
0062
50H
Q00
6345
HQ
0067
61−
HQ
0066
69H
Q00
6580
HQ
0069
52H
Q00
6502
TG
FIN
LA
ND
Noc
tuid
aeE
ustr
oniin
aeD
elto
te u
ncul
aI,
VM
M04
601
GU
8287
03G
U82
8498
GU
8290
26G
U82
9315
GU
8295
79G
U82
9818
GU
8306
91G
U83
0407
GU
8281
92G
U83
0089
FIN
LA
ND
Noc
tuid
aeC
ucul
liina
eC
ucul
lia
umbr
atic
aI,
IV,V
MM
0454
3G
U82
8701
GU
8284
96G
U82
9024
GU
8293
13G
U82
9577
GU
8298
17G
U83
0689
GU
8304
05G
U82
8190
GU
8300
87T
G/T
SFI
NL
AN
D
Noc
tuid
aeD
ilobi
nae
Dil
oba
caer
uleo
ceph
ala
II,V
MM
0926
7JN
4012
46JN
4011
28JN
4013
64JN
4014
75JN
4009
82−
JN40
1887
JN40
1784
−JN
4016
78T
G/T
SFI
NL
AN
D
Noc
tuid
aeR
aphi
inae
Rap
hia
abru
ptI,
II,V
CW
M-9
4-03
72G
U82
8548
GU
8283
50G
U82
8893
GU
8291
93G
U82
9455
GU
8297
28G
U83
0579
GU
8302
70G
U82
8059
GU
8299
39T
GU
SA
Noc
tuid
aePa
nthe
inae
Euc
ocyt
ia m
eeki
II,V
RZ
87JN
4012
47JN
4011
29−
−−
−JN
4018
88JN
4017
85JN
4010
47−
TG
/TS
IND
ON
ES
IA
Noc
tuid
aePa
nthe
inae
Thi
acid
as s
p.II
,VR
Z45
9JN
4012
49JN
4011
31JN
4013
66JN
4014
77JN
4009
37JN
4015
79JN
4018
88JN
4017
87−
JN40
1680
TG
IND
IA
Noc
tuid
aePa
nthe
inae
Ant
itri
sulo
ides
cat
ocal
ina
II,V
RZ
388
JN40
1248
JN40
1130
JN40
1365
JN40
1476
JN40
0936
JN40
1578
JN40
1889
JN40
1786
JN40
1048
JN40
1679
TG
MA
LA
YS
IA
Noc
tuid
aePa
nthe
inae
Pan
thea
coe
nobi
taI,
II,V
MM
0458
3G
U82
8702
GU
8284
97G
U82
9025
GU
8293
14G
U82
9578
−G
U83
0690
GU
8304
06G
U82
8191
GU
8300
88T
G/T
SFI
NL
AN
D
Noc
tuid
aeA
cont
iinae
Em
mel
ia tr
abea
lis
I,V
MM
0989
3H
Q00
6147
HQ
0068
54H
Q00
6245
HQ
0063
40H
Q00
6756
−H
Q00
6664
HQ
0065
76H
Q00
6947
−S
AR
DIN
IA
Noc
tuid
aeA
cont
iinae
Aco
ntia
luci
daI,
IV,V
MM
0015
2G
U82
8617
GU
8284
15G
U82
8952
GU
8292
43G
U82
9512
GU
8297
63G
U83
0627
GU
8303
27G
U82
8118
GU
8300
06T
GG
RE
EC
E
Noc
tuid
aeB
agis
arin
aeA
myn
a oc
toII
,VR
Z50
JN40
1242
JN40
1125
JN40
1360
JN40
1471
JN40
0984
JN40
1574
JN40
1883
JN40
1780
JN40
1043
JN40
1675
MA
LA
YS
IA
Noc
tuid
aeB
agis
arin
aeX
anth
odes
alb
ago
I,II
,VM
M09
894
GU
8288
44G
U92
9808
GU
8291
45G
U82
9412
GU
8296
93−
GU
8307
92G
U83
0535
GU
8283
08G
U83
0224
SA
RD
INIA
Noc
tuid
aeB
agis
arin
aeR
amad
asa
pavo
II,V
RZ
382
JN40
1241
JN40
1124
JN40
1359
JN40
1470
JN40
0978
JN40
1573
JN40
1882
JN40
1779
−JN
4016
74T
SM
AL
AY
SIA
Noc
tuid
aeB
agis
arin
aeD
yrze
la p
lagi
ata
II,V
RZ
395
JN40
1240
JN40
1123
JN40
1358
JN40
1469
JN40
0932
JN40
1572
JN40
1881
JN40
1778
JN40
1042
JN40
1673
TS
MA
LA
YS
IA
Noc
tuid
aeB
agis
arin
aeE
ncru
phio
n le
ena
II,V
RZ
351
JN40
1243
JN
4011
26JN
4013
61JN
4014
72JN
4009
33JN
4015
75JN
4018
84JN
4017
81JN
4010
44JN
4016
76C
OS
TA
RIC
A
Noc
tuid
aeB
agis
arin
aeV
espo
la c
aeru
leif
era
VR
Z59
5X
XX
XX
−X
XX
−T
SC
OS
TA
RIC
A
Noc
tuid
aeB
agis
arin
aeP
aran
giti
a te
mpe
rata
II,I
V,V
RZ
463
JN40
1244
JN40
1127
JN40
1362
JN40
1473
JN40
0934
JN40
1576
JN40
1885
JN40
1782
JN40
1045
−
CO
ST
A R
ICA
Noc
tuid
aeB
agis
arin
aeP
aran
giti
a m
osai
caII
,IV
,VR
Z46
4JN
4012
45−
JN40
1363
JN40
1474
JN40
0935
JN40
1577
JN40
1886
JN40
1783
JN40
1046
JN40
1677
CO
ST
A R
ICA
Noc
tuid
aeB
agis
arin
aeD
iopa
cor
one
II,I
V,V
RZ
472
JN40
1239
JN40
1122
JN40
1357
−JN
4009
31JN
4015
71JN
4018
80JN
4017
77JN
4010
41−
FRE
NC
H
GU
IAN
A
Noc
tuid
aeB
agis
arin
aeC
onca
na m
undi
ssim
aIV
,VR
Z62
1X
XX
XX
XX
−−
XT
SU
SA
Noc
tuid
aeB
agis
arin
aeC
onca
na le
cta
IV,V
RZ
474
X−
−X
X−
XX
−−
CO
ST
A R
ICA
Noc
tuid
aeB
agis
arin
aeC
onca
na p
erm
ixta
IV,V
RZ
482
XX
XX
XX
XX
−X
GU
AT
EM
AL
A
Noc
tuid
aeA
mph
ipyr
inae
Am
phip
yra
perf
lua
I,II
,IV
,VM
M01
162
GU
8286
60G
U82
8458
GU
8289
91G
U82
9275
GU
8295
46G
U82
9787
GU
8306
57
GU
8303
66G
U82
8157
GU
8300
47T
GFI
NL
AN
D
Noc
tuid
aeA
mph
ipyr
inae
Bra
chio
nych
a nu
becu
losa
I,V
MM
0154
2G
U82
8667
GU
8284
65G
U82
8998
GU
8292
81G
U82
9552
GU
8297
93G
U83
0663
GU
8303
73G
U82
8164
GU
8300
54T
SFI
NL
AN
D
Noc
tuid
aeM
etop
oniin
aeF
lam
mon
a qu
adri
fasc
iata
VR
Z59
6X
XX
XX
XX
XX
XT
SM
AL
AY
SIA
Noc
tuid
aeM
etop
oniin
aeP
anem
eria
tene
brat
aI,
VM
M00
005
HQ
0061
57H
Q00
6863
HQ
0062
54H
Q00
6349
HQ
0067
66H
Q00
6437
HQ
0066
73H
Q00
6584
HQ
0069
56H
Q00
6506
FIN
LA
ND
Noc
tuid
aeA
cron
ictin
aeC
rani
opho
ra li
gust
riI,
VM
M06
745
HQ
0061
48H
Q00
6855
HQ
0062
46H
Q00
6341
HQ
0067
57H
Q00
6432
HQ
0066
65H
Q00
6577
HQ
0069
48H
Q00
6498
TS
FIN
LA
ND
Noc
tuid
aeA
cron
ictin
aeA
cron
icta
am
eric
ana
VR
Z59
7X
XX
XX
XX
X−
XT
GU
SA
Noc
tuid
aeA
cron
ictin
aeA
cron
icta
rum
icis
I,V
MM
0152
9G
U82
8666
GU
8284
64G
U82
8997
GU
8292
80G
U82
9551
GU
8297
92G
U83
0662
GU
8303
72G
U82
8163
GU
8300
53T
GFI
NL
AN
D
Noc
tuid
aeA
cron
ictin
aeC
erm
a ce
rint
haV
RZ
617
XX
XX
XX
X−
−X
US
A
Noc
tuid
aeA
cron
ictin
aeC
omac
hara
cad
bury
iV
RZ
618
XX
XX
XX
XX
−X
TS
US
A
Appendix
47
Noc
tuid
aeA
cron
ictin
aeH
arri
sim
emna
tris
igna
taV
RZ
619
XX
XX
XX
XX
−X
US
A
Noc
tuid
aeA
cron
ictin
aeP
olyg
ram
mat
e he
brae
icum
VR
Z62
0X
XX
XX
XX
XX
XT
SU
SA
Noc
tuid
aeA
cron
ictin
aeA
grio
pode
s fa
llax
VR
Z61
6X
XX
XX
XX
X−
XT
SU
SA
Noc
tuid
aeA
gari
stin
aeP
eris
cept
a po
lyst
icta
I,II
,VM
M07
669
GU
8288
20G
U92
9788
GU
8291
25G
U82
9400
GU
8296
74G
U82
9892
GU
8307
73G
U83
0519
GU
8282
89G
U83
0201
TG
/TS
AU
ST
RA
LIA
Noc
tuid
aeA
ediin
aeA
edia
leuc
omel
asII
,VR
Z27
7JN
4012
50JN
4011
32JN
4013
67JN
4014
78JN
4009
76JN
4015
80JN
4018
91JN
4017
88−
JN40
1681
TS
JAP
AN
Noc
tuid
aeC
ondi
cina
eC
ondi
ca il
lect
aV
RZ
511
XX
XX
XX
X−
−X
TG
MA
LA
YS
IA
Noc
tuid
aeC
ondi
cina
eC
ondi
ca v
ecor
sI,
IIC
WM
-95-
0471
GU
8285
50G
U82
8352
GU
8288
95G
U82
9194
GU
8294
57−
GU
8305
81−
GU
8280
61G
U82
9941
TG
US
A
Noc
tuid
aeC
ondi
cina
eH
emic
epha
lis
ales
aII
,VR
Z34
1JN
4012
51JN
4011
33JN
4013
68JN
4014
79JN
4009
38JN
4015
81JN
4018
92JN
4017
89JN
4010
49JN
4016
82C
OS
TA
RIC
A
Noc
tuid
aeH
elio
thin
aeP
yrrh
ia u
mbr
aI,
VM
M05
114
GU
8287
12G
U82
8507
GU
8290
34G
U82
9324
GU
8295
88G
U82
9825
GU
8307
00G
U83
0416
GU
8282
00G
U83
0098
FIN
LA
ND
Noc
tuid
aeun
assi
gned
Chy
toni
x di
ehli
VR
Z51
7X
XX
XX
XX
X−
XM
AL
AY
SIA
Noc
tuid
aeB
ryop
hilin
aeC
ryph
ia r
aptr
icul
aI,
VM
M04
919
GU
8287
08G
U82
8503
GU
8290
31G
U82
9320
GU
8295
84G
U82
9822
GU
8306
96G
U83
0412
GU
8281
96G
U83
0094
FIN
LA
ND
Noc
tuid
aeB
ryop
hilin
aeSt
enol
oba
futi
iV
RZ
523
XX
XX
XX
XX
X−
MA
LA
YS
IA
Noc
tuid
aeun
assi
gned
Ecp
atia
sp.
VR
Z54
5X
XX
XX
XX
X−
XM
AL
AY
SIA
Noc
tuid
aeun
assi
gned
Ecp
atia
long
inqu
uaI,
II,V
RZ
25H
Q00
6190
HQ
0068
94H
Q00
6286
HQ
0063
80H
Q00
6798
−H
Q00
6703
HQ
0066
11H
Q00
6985
HQ
0065
32H
ON
G K
ON
G
Noc
tuid
aeN
octu
inae
Tira
cola
aur
eata
VR
Z14
XX
XX
−X
XX
X−
IND
ON
ES
IA
Noc
tuid
aeN
octu
inae
Act
inot
ia p
olyo
don
I,V
MM
0515
3G
U82
8714
GU
8285
09−
GU
8293
26G
U82
9590
GU
8298
27G
U83
0702
GU
8304
18G
U82
8202
GU
8301
00T
GFI
NL
AN
D
Noc
tuid
aeN
octu
inae
Hop
lodr
ina
octo
gena
ria
I,V
MM
0165
1H
Q00
6153
HQ
0068
60H
Q00
6251
HQ
0063
46H
Q00
6762
HQ
0064
34H
Q00
6670
HQ
0065
81H
Q00
6953
HQ
0065
03FI
NL
AN
D
Noc
tuid
aeN
octu
inae
Dia
phon
e sp
.I,
II,V
MF-
05-0
053
GU
8285
71G
U82
8372
GU
8289
13G
U82
9206
GU
8294
75G
U82
9738
GU
8305
91G
U83
0285
GU
8280
76G
U82
9960
TA
NZ
AN
IA
Noc
tuid
aeN
octu
inae
Apa
mea
cre
nata
I,IV
,VM
M01
170
GU
8286
61G
U82
8459
GU
8289
92G
U82
9276
GU
8295
47G
U82
9788
GU
8306
58G
U83
0367
GU
8281
58G
U83
0048
TG
FIN
LA
ND
Noc
tuid
aeN
octu
inae
Ufe
us fa
unas
I,V
RR
-98-
0914
GU
8288
60G
U92
9822
GU
8291
63G
U82
9425
GU
8297
09G
U82
9911
GU
8308
07G
U83
0552
GU
8283
20G
U83
0238
TG
US
A
Noc
tuid
aeN
octu
inae
Noc
tua
fim
bria
taI,
II,I
V,V
MM
0475
2G
U82
8705
GU
8285
00G
U82
9028
GU
8293
17G
U82
9581
GU
8298
20G
U83
0693
GU
8304
09G
U82
8194
GU
8300
91T
GFI
NL
AN
D