TY - JOUR T1 - Critical re-examination of known purported fossil Bombycoidea (Lepidoptera) JF - PeerJ Y1 - 2023 DO - 10.7717/peerj.16049 A1 - Heikkilä, M. A1 - Minet, J. A1 - Zwick, A. A1 - Hundsdoerfer, A. A1 - Rougerie, R. A1 - Kitching, I.J. SP - e16049 KW - ATTACUS KW - BOMBYCITES KW - BOMBYCOIDEA KW - COCOON KW - FOSSILS KW - MIOCLANIS KW - PROBOSCIS KW - PUPATION CHAMBER KW - SATURNIIDAE KW - SPHINGIDAE KW - SPHINGIDITES AB -

"We critically re-examine 17 records of fossils currently assigned to the lepidopteran superfamily Bombycoidea, which includes the silk moths, emperor moths and hawk moths. These records include subfossils, compression and impression fossils, permineralizations and ichnofossils. We assess whether observable morphological features warrant their confident assignment to the superfamily. None of the examined fossils displays characters that allow unequivocal identification as Sphingidae, but three fossils and a subfossil (Mioclanis shanwangiana Zhang, Sun and Zhang, 1994, two fossil larvae, and a proboscis in asphaltum) have combinations of diagnostic features that support placement in the family. The identification of a fossil pupa as Bunaeini (Saturniidae) is well supported. The other fossils that we evaluate lack definitive bombycoid and, in several cases, even lepidopteran characters. Some of these dubious fossils have been used as calibration points in earlier studies casting doubt on the resulting age estimates. All fossil specimens reliably assigned to Bombycoidea are relatively young, the earliest fossil evidence of the superfamily dating to the middle Miocene."

VL - 11 ER - TY - JOUR T1 - Phylogeny and classification of Endromidae (Lepidoptera: Bombycoidea) based on mitochondrial genomes JF - Arthropod Systematics & Phylogeny Y1 - 2023 A1 - Deng, M. A1 - Zwick, A. A1 - Chen, Q. A1 - Liao, C.-Q. A1 - Wang, W. A1 - Wang, X. A1 - Huang, G.-H. SP - 395 EP - 408 KW - ANDRACA KW - ANDRACINI KW - BAYESIAN INFERENCE KW - COMPARMUSTILIA KW - DALAILAMA KW - ENDROMIDAE KW - ENDROMINAE KW - ENDROMIS KW - FALCOGONA KW - HIGHER CLASSIFICATION KW - MAXIMUM LIKELIHOOD KW - MIRINA KW - MIRININAE KW - MITOGENOME KW - MUSTILIA KW - MUSTILIINI KW - MUSTILIZANS KW - OBERTHUERIA KW - OBERTHUERIINI KW - OBERTHUERINAE KW - PHYLOGENY KW - PRISMOSTICTA KW - PRISMOSTICTINAE KW - PRISMOSTICTOIDES KW - PROMUSTILIA KW - PSEUDANDRACA KW - SESQUILUNA KW - SMERKATA KW - THEOPHOBA AB -

"The small, relict-like moth family Endromidae is well-established within the superfamily Bombycoidea, but relationships within the family have remained vague for the last decade, primarily due to very limited taxon sampling. This resulted in the explicit removal of all internal suprageneric classification by Zwick et al. (2011) when they synonymized Mirinidae and the bombycid subfamilies Oberthueriinae and Prismostictinae with Endromidae. Nucleotide and amino acid data sets of the 13 mitochondrial, protein-coding genes from representatives of 13 of the 16 accepted endromid genera were used to estimate phylogenetic relationships based on maximum likelihood and Bayesian inference methods. The results strongly support Endromidae as a monophyletic group and enable the establishment and diagnosis of four subfamilies (Endrominae, Mirininae stat. rev., Oberthueriinae stat. rev. and Prismostictinae stat. rev.). Within subfamily Oberthueriinae, we establish three tribes: Oberthueriini stat. rev., Andracini tribe nov. and Mustiliini tribe nov. We provide morphological diagnoses and a genus-level checklist for the three tribes. Promustilia yajiangensis Wang, X. & Zolotuhin, 2015 is transferred to Mustilizans as M. yajiangensis comb. nov. to establish reciprocal monophyly of the two genera, and Andraca gongshanensis is transferred to Pseudandraca as P. gongshanensis comb. nov. We also synonymize Andraca (Chrypathemola) syn. nov. with Andraca (Andraca), as the latter is deeply nested within the former."

VL - 81 UR - https://doi.org/10.3897/asp.81.e90721 ER - TY - JOUR T1 - The principal structure of male genital sclerites and muscles of bombycoid moths, with special reference to Anthelidae (Lepidoptera: Bombycoidea) JF - Arthropod Structure & Development Y1 - 2009 DO - 10.1016/j.asd.2008.07.006 A1 - Zwick, A. SP - 147–161 KW - ANTHELIDAE KW - BOMBYCOIDEA KW - FUNCTION KW - GENITAL MUSCULATURE KW - GNATHOS KW - HOMOLOGY KW - JUXTA KW - MALE GENITALIA KW - PHALLUS KW - SOCII KW - SUBSCAPHIUM KW - TRANSTILLA KW - UNCUS KW - VALVE AB -

"Male genital structures and muscles of bombycoid moths have repeatedly been misidentified in the literature. Furthermore, the genital structures of some bombycoid families, such as the poorly known Australo-New Guinean Anthelidae, have essentially remained unstudied. Based on comparative morphology, this study details the principal arrangements of male genital sclerites and muscles in all bombycoid families, with particular focus on basic structures and their modifications in Anthelidae. Emphasis is placed on the homology of and fusions between these structures and their function, providing a basis for the interpretation of modifications in future phylogenetic and taxonomic studies. This includes the unique fusion of gnathos and valvae in several bombycoid families, the arrangement and extent of the fused tegumen and vinculum, as well as the homology of the ‘‘transtilla’’. Further, a modification of the valve adductor muscle (the segment IX sternum to valva muscle, m4) widely regarded as a synapomorphy of Bombycoidea is demonstrated to be non-existent, as is the presumed presence of the valve abductor muscle (the segment IX tergum to valva muscle, m2) in Saturniidae."

VL - 38 ER - TY - JOUR T1 - Präimaginalstadien und Biologie von Syrastrena dorca Zolotuhin, 2005 (Lepidoptera: Lasiocampidae) JF - Entomologische Zeitschrift Y1 - 2020 A1 - Ihle, S. A1 - Zwick, A. SP - 217 EP - 221 KW - CHINA KW - FOODPLANT KW - IMMATURE STAGES KW - LASIOCAMPIDAE KW - MALE GENITALIA KW - PUPA KW - SYRASTRENA VL - 130 IS - 4 ER - TY - JOUR T1 - Phylogenomics resolves major relationships and reveals significant diversification rate shifts in the evolution of silk moths and relatives. JF - BMC Evolutionary Biology Y1 - 2019 DO - 10.1186/s12862-019-1505-1 A1 - Hamilton, C.A. A1 - St Laurent, R.A. A1 - Dexter, K. A1 - Kitching, I.J. A1 - Breinholt, J.W. A1 - Zwick, A. A1 - Timmermans, M.J.T.N. A1 - Barber, J.R. A1 - Kawahara, A.Y. SP - 182 [1 EP - 13] KW - ANCHORED HYBRID ENRICHMENT KW - APATELODIDAE KW - AROTROS KW - BOMBYCIDAE KW - BOMBYCOIDEA KW - DNA KW - PHYLOGENOMICS KW - PHYLOGENY KW - TARGETED SEQUENCE CAPTURE AB -

"Background: Silkmoths and their relatives constitute the ecologically and taxonomically diverse superfamily Bombycoidea, which includes some of the most charismatic species of Lepidoptera. Despite displaying spectacular forms and diverse ecological traits, relatively little attention has been given to understanding their evolution and drivers of their diversity. To begin to address this problem, we created a new Bombycoidea-specific Anchored Hybrid Enrichment (AHE) probe set and sampled up to 571 loci for 117 taxa across all major lineages of the Bombycoidea, with a newly developed DNA extraction protocol that allows Lepidoptera specimens to be readily sequenced from pinned natural history collections.
Results: The well-supported tree was overall consistent with prior morphological and molecular studies, although some taxa were misplaced. The bombycid Arotros Schaus was formally transferred to Apatelodidae. We identified important evolutionary patterns (e.g., morphology, biogeography, and differences in speciation and extinction), and our analysis of diversification rates highlights the stark increases that exist within the Sphingidae (hawkmoths) and Saturniidae (wild silkmoths).
Conclusions: Our study establishes a backbone for future evolutionary, comparative, and taxonomic studies of Bombycoidea. We postulate that the rate shifts identified are due to the well-documented bat-moth “arms race”. Our research highlights the flexibility of AHE to generate genomic data from a wide range of museum specimens, both age and preservation method, and will allow researchers to tap into the wealth of biological data residing in natural history collections around the globe."

VL - 19 UR - https://doi.org/10.1186/s12862-019-1505-1 ER - TY - JOUR T1 - Phylogenetic relationships among Bombycidae s.l. (Lepidoptera) based on analyses of complete mitochondrial genomes JF - Systematic Entomology Y1 - 2019 DO - 10.1111/syen.12337 A1 - Wang, X. A1 - Chen, Z.-M. A1 - Gu, X.-S. A1 - Wang, M. A1 - Huang, G.-H. A1 - Zwick, A. SP - 490 EP - 498 KW - ANDRACA KW - BAYESIAN INFERENCE KW - BOMBYCIDAE KW - BOMBYX KW - COMPARMUSTILIA KW - DNA KW - ENDROMIDAE KW - ERNOLATIA KW - GANISA KW - MAXIMUM LIKELIHOOD KW - MITOGENOME KW - MUSTILIA KW - MUSTILIZANS KW - OCINARA KW - PHYLOGENY KW - PRISMOSTICTA KW - PRISMOSTICTOIDES AB -

"The family Bombycidae (sensu Minet, 1994) is a diverse group of species belonging to the superfamily Bombycoidea. It is an economically important group of moth species, containing well-known silk-producing insects, as well as many pests of agriculture and forestry. The morphology-based hypothesis of Minet (1994) on the composition of Bombycidae is in conflict with subsequent phylogenetic hypotheses for the superfamily based on nuclear genes. In this paper, the complete mitochondrial genomes of nine species of Bombycidae are presented for the first time. Based on these genomes, four dataset partitions and three gblocks parameter settings, phylogenetic relationships among Bombycidae were reconstructed using maximum likelihood and Bayesian inference methods. Bombycidae was confirmed as a polyphyletic group, with the traditional subfamilies Prismostictinae and Oberthueriinae forming a single well-supported clade that is distant to Bombycinae. The phylogenetic relationships within Bombycoidea were supported as ((((Bombycinae, Sphingidae), Saturniidae), (Prismostictinae, Oberthueriinae)), Eupterotidae)."

VL - 44 UR - https://doi.org/10.1111/syen.12337 IS - 3 ER - TY - JOUR T1 - A global checklist of the Bombycoidea (Insecta: Lepidoptera) JF - Biodiversity Data Journal Y1 - 2018 DO - 10.3897/BDJ.6.e22236 A1 - Kitching, I.J. A1 - Rougerie, R. A1 - Zwick, A. A1 - Hamilton, C.A. A1 - St._Laurent, R. A1 - Naumann, S. A1 - Ballesteros-Mejia, L. A1 - Kawahara, A.Y. SP - e22236 KW - ANTHELIDAE KW - APATELODIDAE KW - BOMBYCIDAE KW - BRAHMAEIDAE KW - CARTHAEIDAE KW - CATALOGUE KW - CHECKLIST KW - ENDROMIDAE KW - EUPTEROTIDAE KW - HIGHER CLASSIFICATION KW - ORECTA KW - PHIDITIIDAE KW - SATURNIIDAE KW - SPHINGIDAE AB -

"Background
Bombycoidea is an ecologically diverse and speciose superfamily of Lepidoptera. The superfamily includes many model organisms, but the taxonomy and classification of the superfamily has remained largely in disarray. Here we present a global checklist of Bombycoidea. Following Zwick (2008) and Zwick et al. (2011), ten families are recognized: Anthelidae, Apatelodidae, Bombycidae, Brahmaeidae, Carthaeidae, Endromidae, Eupterotidae, Phiditiidae, Saturniidae and Sphingidae. The former families Lemoniidae and Mirinidae are included within Brahmaeidae and Endromidae respectively. The former bombycid subfamilies Oberthueriinae and Prismostictinae are also treated as synonyms of Endromidae, and the former bombycine subfamilies Apatelodinae and Phitditiinae are treated as families.


New information
This checklist represents the first effort to synthesize the current taxonomic treatment of the entire superfamily. It includes 12,159 names and references to their authors, and it accounts for the recent burst in species and subspecies descriptions within family Saturniidae (ca. 1,500 within the past 10 years) and to a lesser extent in Sphingidae (ca. 250 species over the same period). The changes to the higher classification of Saturniidae proposed by Nässig et al. (2015) are rejected as premature and unnecessary. The new tribes, subtribes and genera described by Cooper (2002) are here treated as junior synonyms. We also present a new higher classification of Sphingidae, based on Kawahara et al. (2009), Barber and Kawahara (2013) and a more recent phylogenomic study by Breinholt et al. (2017), as well as a reviewed genus and species level classification, as documented by Kitching (2018)."

VL - 6 UR - https://doi.org/10.3897/BDJ.6.e22236 ER - TY - JOUR T1 - Increased gene sampling yields robust support for higher-level clades within Bombycoidea (Lepidoptera) JF - Systematic Entomology Y1 - 2011 DO - 10.1111/j.1365-3113.2010.00543.x A1 - Zwick, A. A1 - Regier, J.C. A1 - Mitter, C. A1 - Cummings, M.P. SP - 31 EP - 43 KW - BOMBYCIDAE KW - BOMBYCOIDEA KW - CARTHAEIDAE KW - DNA KW - ENDROMIDAE KW - EPIINAE KW - HIGHER CLASSIFICATION KW - MAXIMUM LIKELIHOOD KW - MIRINIDAE KW - OBERTHUERINAE KW - PHIDITIIDAE KW - PHYLOGENY KW - PRISMOSTICTINAE AB -

"This study has as its primary aim the robust resolution of higher-level relationships within the lepidopteran superfamily Bombycoidea. Our study builds on an earlier analysis of five genes (∼6.6 kbp) sequenced for 50 taxa from Bombycoidea and its sister group Lasiocampidae, plus representatives of other macrolepidoteran superfamilies. The earlier study failed to yield strong support for the monophyly of and basal splits within Bombycoidea, among others. Therefore, in an effort to increase support specifically for higher-level nodes, we generated 11.7 kbp of additional data from 20 genes for 24 of 50 bombycoid and lasiocampid taxa. The data from the genes are all derived from protein-coding nuclear genes previously used to resolve other lepidopteran relationships. With these additional data, all but a few higher-level nodes are strongly supported. Given our decision to minimize project costs by augmenting genes for only 24 of the 50 taxa, we explored whether the resulting pattern of missing data in the combined-gene matrix introduced a nonphylogenetic bias, a possibility reported by others. This was achieved by comparing node support values (i.e. nonparametric bootstrap values) based on likelihood and parsimony analyses of three datasets that differ in their number of taxa and level of missing data: 50 taxa/5 genes (dataset A), 50 taxa/25 genes (dataset B) and 24 taxa/25 genes (dataset C). Whereas datasets B and C provided similar results for common nodes, both frequently yielded higher node support relative to dataset A, arguing that: (i) more data yield increased node support and (ii) partial gene augmentation does not introduce an obvious nonphylogenetic bias. A comparison of single-gene bootstrap analyses identified four nodes for which one or two of the 25 genes provided modest to strong support for a grouping not recovered by the combined-gene result. As a summary proposal, two of these four groupings (one each within Bombycoidea and Lasiocampidae) were deemed sufficiently problematic to regard them as unresolved trichotomies. Since the alternative groupings were always highly localized on the tree, we did not judge a combined-gene analysis to present a problem outside those regions. Based on our robustly resolved results, we have revised the classification of Bombycoidea: the family Bombycidae is restricted to its nominate subfamily, and its tribe Epiini is elevated to subfamily rank (Epiinae stat.rev.), whereas the bombycid subfamily Phiditiinae is reinstated as a separate family (Phiditiidae stat.rev.). The bombycid subfamilies Oberthueriinae Kuznetzov & Stekolnikov, 1985, syn.nov. and Prismostictinae Forbes, 1955, syn.nov., and the family Mirinidae Kozlov, 1985, syn.nov. are established as subjective junior synonyms of Endromidae Boisduval, 1828. The family Anthelidae (Lasiocampoidea) is reincluded in the superfamily Bombycoidea."

VL - 36 IS - 1 ER - TY - JOUR T1 - A large-scale, higher-Level, molecular phylogenetic study of the insect order Lepidoptera (moths and butterflies) JF - PLoS ONE Y1 - 2013 DO - 10.1371/journal.pone.0058568 A1 - Regier, J.C. A1 - Mitter, C. A1 - Zwick, A. A1 - Bazinet, A.L. A1 - Cumming, M.P. A1 - Kawahara, A.Y. A1 - Sohn, J.-C. A1 - Zwickl, D.J. A1 - Cho, S. A1 - Davis, D.R. A1 - Baixeras, J. A1 - Brown, J. A1 - Parr, C. A1 - Weller, S. A1 - Lees, D.C. A1 - Mitter, K.T. SP - e58568 KW - 109FIN KW - 265FIN KW - 268FIN KW - 3007FIN KW - ACC KW - APODITRYSIA KW - BARCODING KW - BOOTSTRAP KW - CAD KW - COI KW - DDC KW - DNA KW - ENOLASE KW - GARLI KW - HIGHER CLASSIFICATION KW - LEPIDOPTERA KW - MACROHETEROCERA KW - MAXIMUM LIKELIHOOD KW - MTDNA KW - PHYLOGENY KW - ROGUE TAXA KW - TINEOIDEA AB -

"Background: Higher-level relationships within the Lepidoptera, and particularly within the species-rich subclade Ditrysia, are
generally not well understood, although recent studies have yielded progress. We present the most comprehensive
molecular analysis of lepidopteran phylogeny to date, focusing on relationships among superfamilies.

Methodology / Principal Findings: 483 taxa spanning 115 of 124 families were sampled for 19 protein-coding nuclear
genes, from which maximum likelihood tree estimates and bootstrap percentages were obtained using GARLI. Assessment
of heuristic search effectiveness showed that better trees and higher bootstrap percentages probably remain to be
discovered even after 1000 or more search replicates, but further search proved impractical even with grid computing.
Other analyses explored the effects of sampling nonsynonymous change only versus partitioned and unpartitioned total
nucleotide change; deletion of rogue taxa; and compositional heterogeneity. Relationships among the non-ditrysian
lineages previously inferred from morphology were largely confirmed, plus some new ones, with strong support. Robust
support was also found for divergences among non-apoditrysian lineages of Ditrysia, but only rarely so within Apoditrysia.
Paraphyly for Tineoidea is strongly supported by analysis of nonsynonymous-only signal; conflicting, strong support for
tineoid monophyly when synonymous signal was added back is shown to result from compositional heterogeneity.

Conclusions / Significance: Support for among-superfamily relationships outside the Apoditrysia is now generally strong.
Comparable support is mostly lacking within Apoditrysia, but dramatically increased bootstrap percentages for some nodes
after rogue taxon removal, and concordance with other evidence, strongly suggest that our picture of apoditrysian
phylogeny is approximately correct. This study highlights the challenge of finding optimal topologies when analyzing
hundreds of taxa. It also shows that some nodes get strong support only when analysis is restricted to nonsynonymous
change, while total change is necessary for strong support of others. Thus, multiple types of analyses will be necessary to
fully resolve lepidopteran phylogeny."

VL - 8 IS - 3 ER - TY - JOUR T1 - Molecular phylogeny of Anthelidae and other bombycoid taxa (Lepidoptera: Bombycoidea) JF - Systematic Entomology Y1 - 2008 DO - 10.1111/j.1365-3113.2007.00410.x A1 - Zwick, A. SP - 190 EP - 209 KW - ANTHELA KW - ANTHELIDAE; APATELODIDAE KW - BOMBYCIDAE KW - BOMBYCOIDEA KW - BRAHMAEIDAE KW - CAD KW - CLASSIFICATION KW - DNA KW - EF-1ALPHA KW - FOODPLANT EVOLUTION KW - LEMONIIDAE KW - PHYLOGENY AB -

"Based on DNA sequences of the fusion protein carbamoylphosphate synthetase/aspartate transcarbamylase/dihydroorotase (CAD; 680 bp) and elongation factor-1α (Ef-1α; 1240 bp); the first molecular phylogeny of the moth family Anthelidae and its placement within the Bombycoidea sensu Brock (1971) (= bombycoid complex sensu Minet, 1994) is proposed. The results strongly support the monophyly of the family Anthelidae and its subfamilies Munychryiinae and Anthelinae, but demonstrate the vast polyphyly of its main genus Anthela Walker, 1855. The proposed phylogeny suggests that grass feeding, as apparent from some pest records, probably is an ancestral trait within the subfamily Anthelinae. Evolutionary relationships of the family Anthelidae and of most parts of the Bombycoidea remain obscure. However, the results contradict many of the widely accepted phylogenetic hypotheses within the Bombycoidea proposed by Minet (1994: Entomologica scandinavica, 25, 63–88). The Brahmaeidae are paraphyletic relative to the Lemoniidae (syn.nov.), and the current concept of Bombycidae is polyphyletic, with the bombycid subfamily Apatelodinae being part of a monophylum comprising Brahmaeidae / Lemoniidae, Eupterotidae and Apatelodidae (stat.rev.)."

VL - 33 IS - 1 ER - TY - JOUR T1 - Notes on the hawkmoths of the Philippines (Lepidoptera: Sphingidae) JF - Nachrichten des Entomologischen Vereins Apollo (N.F.) Y1 - 2001 A1 - Zwick, A. A1 - Treadaway, C.G. SP - 177 EP - 181 KW - CATALOGUE KW - MACROGLOSSUM KW - MALE GENITALIA KW - PALAWAN KW - PHILIPPINES KW - SPHINGIDAE KW - WORLDMAP1 VL - 22 UR - https://www.zobodat.at/publikation_volumes.php?id=50774 N1 - N.F. ER -