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Research Project: Systematics of Moths Significant to Biodiversity, Quarantine, and Control, with a Focus on Invasive Species

Location: Systematic Entomology Laboratory

2020 Annual Report

The order Lepidoptera, specifically moths, represents one of the greatest radiations of herbivorous animals on the planet. The research component of this plan focuses on the most economically important groups of moths. Gaps in our knowledge of morphological, biological, and molecular characters, and the paucity of phylogenetic analyses, within many groups in these families combine to slow progress on the development of identification tools for use in pest detection, exclusion, and management, and in the successful implementation of biological control projects. The primary focus of this plan is to remedy some of those shortcomings by making available to a broad audience tools, databases, and images that will facilitate identifications and research over a wide range of economically important groups of moths. Within the Lepidoptera specific taxonomic groups are selected for revision based on specific expertise, the need for revisionary work, and the relevance of the group to American agriculture. This project also includes a service component that draws upon SEL experts to identify specimens for regulatory agencies (often on an urgent basis) and other research agencies and stakeholders, and to maintain and enhance portions of the National Insect Collection. Over the next five years, we will be addressing the following objectives: (1) conduct integrative taxonomic research that wherever possible incorporates larval and molecular data, in addition to adult data, to determine and circumscribe species and manage associated information, as follows: recognize and describe new and/or cryptic species; develop identification keys and illustrations; refine hypotheses of relationships to be reflected in classification; track distributions, and investigate regional host use and specificity of moths that are pests, potential pests, invasive species, and/or species beneficial to U.S. agriculture of Pyraloidea, Gelechiinae, Noctuoidea, and grass-feeding patterns in moths; (2) manage and enhance via fieldwork appropriate segments of the U.S. National Insect Collection to enable morphological and molecular research, mine the associated distributional and biological data for comprehensive databases, and provide identifications as needed; and (3) provide expert/authoritative identifications and generate research associated with specimens submitted by ARS researchers and other stakeholders or intercepted at U.S. ports by APHIS, Homeland Security, and state departments of agriculture for early detection of potentially invasive or novel pests.

This project will undertake research on a number of economically important plant feeding moths. We will generate morphological, molecular (DNA sequences), and biological characters that will be used to test species concepts and hypotheses of relationship among agriculturally important moths. These data also will be used to develop new diagnostic tools (descriptions, images, illustrations, keys) to permit more rapid and accurate identifications. Databases containing scientific names, distributions, taxonomic literature, and host plant and specimen data pertaining to economically important moths will be expanded and disseminated to the user community. These and other taxonomic tools will be made accessible to the public via publications, the internet, and other electronic media. Timely and accurate identifications of moths will be provided, including those intercepted at ports-of-entry by APHIS-PPQ or submitted by a wide range of scientists and regulatory agencies. Portions of the National Insect Collection at the National Museum of Natural History, a vital tool for research and identification, will be maintained and enhanced via fieldwork.

Progress Report
This is the final report for project 8042-22000-294-00D, Systematics of Moths Significant to Biodiversity, Quarantine, and Control, with a Focus on Invasive Species, which will terminate in December 2020. The new project plan finishing up NP304 OSQR Review. Substantial results were realized over the 3.5-year project that began December 2016 and terminated early to synchronize with other projects in the laboratory. The writing and evaluation of the new project is currently underway. Progress was made on all three objectives, which fall under National Program 304, Crop Protection and Quarantine, Component 1, Systematics and Identification. Particularly notable progress was made on research targeting Problem Statement 1A, the identification of pests or potential pests of the Nation’s crops and natural ecosystems, as well as exotic insects that could be used as biological control agents or show potential as biological control agents of invasive plants. Significant progress was made in the main research Objective 1, which was to conduct integrative taxonomic research within three, speciose and economically important moth groups, the Pyraloidea, Gelechiinae, and Noctuoidea. Sixteen specific moth taxa were targeted. Combinations of adult, larval, and molecular data were used to diagnose and circumscribe species, revise their classification, communicate associated information, and explore questions relevant to the emergence of major and/or invasive pests. In addition to our focal taxa, we discovered, described, and classified new moth species whose larvae feed on a wide variety of commodities and economically important plants, including legumes, pineapples, corn, sorghum, sugarcane, nightshade family, and including ecologically important, native, North American grasses. Value-added imaging and molecular research included significant progress to elucidate the complex genitalia morphology of gelechiid moths through the use of 3D imaging, and the assembly and annotation of transcriptomes of two species of the noctuoid group Papaipema, one a fern-feeder and another a grass-feeder, which contributed to our fourth research subobjective to circumscribe and better understand focal groups of grass-feeding specialists. We made substantial progress on our fourth subobjective by completing the mining of three data sets, identifying preliminary patterns and formulating testable hypotheses, refining our data queries and circumscriptions of grass-feeding focal taxa to test specific patterns. Our extensive data mining efforts produced taxonomic profiles of diets based on over 350,000 moth interceptions at U.S. ports to explore patterns and trends over 25+ years. Successful studies of pests, and/or potential pests, included the use of molecular tools for the identification of the Old World Bollworm, Helicoverpa armigera, intercepted at U.S. ports. The latter study also demonstrated the unreliability of host and origin data for screening purposes. Studies on sugarcane borers that are major pests in the Western Hemisphere of food crops, such as corn, rice, and wheat, led to the discovery of a new, cryptic species feeding on Eastern gamagrass in the United States, and clarified the identity and distribution of other sugarcane-feeding species in Colombia. Moths and their larvae exhibit great host and habitat diversity and, as part of our general objective, we also studied and published results on moths that feed on orchids and ferns, moths that live in bird nests, maple-feeding leaf miners, and a new host plant family for the destructive navel orangeworm. Our research also included biology and identify of aquatic caterpillars, including two chapters in aquatic insect books, one of which is used by university students in the United States, and description of aquatic species new to the United States. Aquatic caterpillars feed on native and invasive aquatic weeds that are important to the health of U.S. waterways and detection for quarantine purposes. Our mandate called for collaboration with specialists in our field due to the nature of invasive species from other areas worldwide. Significant findings included the discovery, in collaboration with ARS biological control laboratories, of new species feeding on the Brazilian peppertree, one of the most aggressive invasive weeds in the United States, and the Old World Climbing fern-feeders in the Everglades. We also collaborated with multiple international teams on projects focused on the evolution of economically significant moth groups. These included a global review of pyraloid fossils, and a study of the Old World stem-borers which are primary pests of crop grasses. The latter work combined paleoenvironmental proxies with molecular phylogenetic data to demonstrate an inverse correlation between the diversification of the larvae and that of grasses as a function of temperature and atmospheric composition. Additionally, significant progress in most projects was derived from a collaboration with, and externally funded by, Smithsonian Institution and the Biodiversity Institute of Ontario and Barcode of Life Data Systems, known as BOLD, to generate DNA COI barcodes for all North American moth species. Specimens were prioritized, prepared, and submitted for sequencing, many of them old or very old specimens that required next-generation sequencing techniques. The sequencing success rate for the ancient DNA was over 80%. In the course of this work, protocols for dissections of National Museum of Natural History Noctuoidea type specimens were implemented to include mandatory DNA extractions. Two of our objectives were service components linked to research Objective 1. The first was to maintain and enhance portions of the National Museum of Natural History, Smithsonian Institution collection. Our research results were embedded and vouchered in our curatorial work. A major focus and basis of taxonomic research included imaging, morphological, and molecular study of moth type specimens at the National Museum of Natural History. Significant progress was made in the imaging of gelechioid and noctuoid type specimens. We verified and added over 2,000 additional photographs of gelechioid moth type specimens, their labels, and slide dissections, to the Lepidoptera Specimen Database of the National Museum of Natural History. This resource is crucial to resolve identities of economically important moths. The oldest and largest U.S. collection of a snout moth grass-feeding group, which includes the sod webworm, was curated and databased to mine biological information about host use for our fourth objective on grass-feeding specialists. The second objective, to identify potential pest moths, has direct, daily and long-term impact. We identified specimens for regulatory agencies, primarily the Animal and Plant Health Inspection Service, and other research agencies and stakeholders. During the life of this project we made 10,379 identifications of 10,833 specimens. This included 6,367 URGENT identifications (i.e., to be done immediately as they are submitted by U.S. port identifiers for potential quarantine action on imported commodities). Routine submissions were from local agricultural and environmental agencies, Animal and Plant Health Inspection Service identifiers to verify training efficacy and status of expertise, and from U.S. citizens. The overall impact of the research conducted in this project is the discovery of new moth species; hypotheses of their evolution; and their biological, evolutionary, and economic relationships to American agriculture and natural resources. Significant direct impact was made when we provided identifications, tools, databases, and images to facilitate identifications of economically important moth taxa for use in pest detection at U.S. ports, exclusion, and management, and in the successful implementation of biological control projects.

1. New twirler moth genera and species to combat the Brazilian Peppertree. Peppertree is one of the most aggressive invasive weeds in Florida and other parts of its invaded range. An ARS researcher located at the National Museum of Natural History, Smithsonian Institution, Washington, D.C., hit both marks of research and intellectual sharing by collaborating with ARS scientists in the Invasive Plant Research Lab in Fort Lauderdale, Florida, in their efforts to find insect agents for biocontrol of Brazilian Peppertree. Together, they discovered and described five new species in three new genera of twirler moths from the invasive plant's native region in South America. This research provided a better perspective on the diversity of herbivores, how they are related, and host range across a twirler moth subfamily.

2. Discovery of A “Missing Link” Twirler moth. Twirler moths are represented by thousands of species and numerous agricultural and forest pests. Understanding how twirler moth species from different parts of the World are related and where they occur facilitate research to mediate their impact on U.S. natural resources. An ARS researcher located at the National Museum of Natural History, Smithsonian Institution, Washington, D.C., mentoring a doctoral graduate student at the University of Panama, discovered and described a new, missing link" species of twirler moth. The new species is from Panama and related to a poorly understood species described in 1923 from Brazil based on only male specimens. Prior to this research, the genealogy of the 100-year-old-moth was unknown. Discovery of the new moth species from Panama and the morphology of the female allowed a hypothesis of the relatedness of both species to other twirler moths. The results of this research help us understand broader evolutionary relationships, diversity, and geographic distribution among the World's twirler moths.

3. Tissue acquisition for the taxonomic resolution of the fall armyworm. An ARS scientist located at the National Museum of Natural History, Smithsonian Institution, Washington, D.C., collaborated with a scientist at the Identification Technology Program Molecular Laboratory, Animal and Plant Health Inspection Service, Fort Collins, Colorado, to procure tissue from type specimens in foreign museums of synonyms. The Fall Armyworm is one of the World's most high-profile economically important pests, which has invaded and become established in Africa, Asia, and Australia in the last four years. The purpose of this work is to correct an improper designation of a neotype, which impedes accurate scientific communication concerning this species. This work is being combined with ongoing molecular diagnostic data from intercepted specimens to characterize the many pest species of Spodoptera.

4. New species and genera of fern-feeding Noctuidae. An ARS scientist located at the National Museum of Natural History, Smithsonian Institution, Washington, D.C., collaborated with a colleague at The Natural History Museum, London, to revise and describe new taxa of fern-feeding Noctuidae from Central and South America. This research is pivotal to understanding the geographic and phylogenetic origins of fern diets considered rare in nature, and more immediately to the classification of the subfamily to which they belong.

5. Analytical applications and trends in DNA barcoding. An ARS scientist located at the National Museum of Natural History, Smithsonian Institution, Washington, D.C., collaborated with a scientist at the American Museum of Natural History, New York, to examine over 3,750 publications that utilized DNA barcoding, and evaluate trends in empirical technique, applications, and interpretations of analyses. The paper described a sharp rise in publications using DNA barcoding for forensic and conservation research in 2017, and noted that despite the rapid expansion of DNA barcode-based research driven largely by the discovery and description of new species, analytical misapplications remain common.

Review Publications
Stonis, J.R., Remeikis, A., Diskus, A., Orlovskyte, S., Vargas, S., Solis, M.A. 2019. A new leaf-mining pest of guava, Hesperolyra guajavifoliae sp. n., with comments on the diagnostics of the endemic Neotropical genus Hesperolyra van Nieukerken (Lepidoptera, Nepticulidae). ZooKeys. 900:87-110. http;10.3897/zookeys.900.46332.
Corro Chang, P., Metz, M. 2020. Universidad de Panamá, Vicerrectoría de Investigación y Postgrado. Doctorado en Ciencias Biológicas con énfasis en Entomología, Panamá; Agricultural Research Service - Systematic Entomology Lab. Proceedings of the Entomological Society of Washington. 122(2):291-298.
Desalle, R., Goldstein, P.Z. 2019. Interpretive trends in DNA barcoding. Frontiers in Ecology and Evolution. 7(302).
Dolibaina, D., Casagrande, M., Specht, A., Mielke, O.N., Legrain, A., Zilli, A., Goldstein, P.Z. 2019. Taxonomy of the rivorum species-group of Leucania Ochsenheimer, 1816 (Lepidoptera: Noctuidae). Zootaxa. 4711(3):545-560.
Goldstein, P.Z., Zilli, A. 2019. Thraumata, a new genus from South America with a description of a new species from Peru (Lepidoptera: Noctuidae: Eriopinae). ZooKeys. 866:139-160.
Goldstein, P.Z., Janzen, D.H., Hallwachs, W. 2019. Aprica: A new genus and life history for the pteridivore "Xanthia patula Druce, 1898. ZooKeys. 866:127-145.
Mccarty, M.E., Adamski, D., Metz, M., Landry, J.F. 2020. Two new species in the genus Frumenta Busck 1939 (Lepidoptera: Gelechiidae: Gnorimoschemini) with discovery of a culcitula in the male. Proceedings of the Entomological Society of Washington. 122(2):415-441.
Metz, M., Wheeler, G.S., Mckay, F., Dyer, K.G. 2019. New genera and species of Gelechiinae (Lepidoptera: Gelechiidae) from South America feeding on Brazilian peppertree. Proceedings of the Entomological Society of Washington. 121(3):461-483.
Solis, M.A. 2019. Aquatic and semiaquatic Lepidoptera In: Aquatic Insects of North America, R. W. Merritt, K.W. Cummins, and M.B. Berg (Eds.). 5th edition. Book Chapter. 765-789.
Solis, M.A., Hayden, J.E., Vargas, S.F., Gonzalez, F., Sanabria, U.C., Gulbronson, C. 2019. A new pyraloid moth species of Sufetula Walker (Crambidae) feeding on pineapple, Ananas comosus (L.) (Bromeliaceae) from Costa Rica. Proceedings of the Entomological Society of Washington. 121(3):497-510.
Solis, M.A., Osoria-Mejia, P., Sarmiento-Naizaque, Z., Barreto-Triana, N. 2020. A new species of Eoreuma Ely (Crambidae: Crambinae) feeding on sugarcane from Colombia. Proceedings of the Entomological Society of Washington. 122(2):471-481.
Solis, M.A., Philips-Rodriguez, E., Hallwachs, W., Dapkey, T., Janzen, D.H. 2020. Asturodes Amsel (Lepidoptera: Crambidae: Spilomelinae): Three new species from the Western Hemisphere and foodplant records from Area de Conservación Guanacaste, Costa Rica. Proceedings of the Entomological Society of Washington. 122(1):147-171.
Stonis, J.R., Diskus, A., Remeikis, A., Vargas, S., Solis, M.A. 2020. Diagnostics and updated catalogue of Acalyptris Meyrick, the second largest genus of Nepticulidae (Lepidoptera) in the Americas. Zootaxa. 4748(2):201-247.
Stonis, J.R., Remeikis, A., Diskus, A., Davis, D.R., Solis, M.A. 2020. American Tischeriidae (Lepidoptera) species from the collection of the National Museum of Natural History, Washington, D.C. Proceedings of the Entomological Society of Washington. 122(3):485-505.
Wheeler, G.S., Dyer, K.G., Metz, M. 2019. Host range of the leaf-tier Tentamen atrivirgulatum (Lepidoptera: Gelechiidae); an unsuitable candidate for biological control of Brazilian peppertree. Biocontrol Science and Technology. 29(12):1172-1180.