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Coleoptera: Lampyridae)
Brigham Young University BYU ScholarsArchive Theses and Dissertations 2020-03-23 Advances in the Systematics and Evolutionary Understanding of Fireflies (Coleoptera: Lampyridae) Gavin Jon Martin Brigham Young University Follow this and additional works at: https://scholarsarchive.byu.edu/etd Part of the Life Sciences Commons BYU ScholarsArchive Citation Martin, Gavin Jon, "Advances in the Systematics and Evolutionary Understanding of Fireflies (Coleoptera: Lampyridae)" (2020). Theses and Dissertations. 8895. https://scholarsarchive.byu.edu/etd/8895 This Dissertation is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected]. Advances in the Systematics and Evolutionary Understanding of Fireflies (Coleoptera: Lampyridae) Gavin Jon Martin A dissertation submitted to the faculty of Brigham Young University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Seth M. Bybee, Chair Marc A. Branham Jamie L. Jensen Kathrin F. Stanger-Hall Michael F. Whiting Department of Biology Brigham Young University Copyright © 2020 Gavin Jon Martin All Rights Reserved ABSTRACT Advances in the Systematics and Evolutionary Understanding of Fireflies (Coleoptera: Lampyridae) Gavin Jon Martin Department of Biology, BYU Doctor of Philosophy Fireflies are a cosmopolitan group of bioluminescent beetles classified in the family Lampyridae. The first catalogue of Lampyridae was published in 1907 and since that time, the classification and systematics of fireflies have been in flux. Several more recent catalogues and classification schemes have been published, but rarely have they taken phylogenetic history into account. Here I infer the first large scale anchored hybrid enrichment phylogeny for the fireflies and use this phylogeny as a backbone to inform classification. -
Coleoptera: Lampyridae, Lampyrinae, Photinini) Endemic to the Brazilian Atlantic Rainforest, with Description of Three New Species
Zootaxa 3835 (3): 325–337 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2014 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3835.3.2 http://zoobank.org/urn:lsid:zoobank.org:pub:C8338F48-3E18-477D-A134-11778ABD6781 Ybytyramoan, a new genus of fireflies (Coleoptera: Lampyridae, Lampyrinae, Photinini) endemic to the Brazilian Atlantic Rainforest, with description of three new species LUIZ FELIPE LIMA DA SILVEIRA1,2 & JOSÉ RICARDO M. MERMUDES1 1Laboratório de Entomologia, Departmento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, A1-107, Bloco A, Av. Carlos Chagas Filho, 373, Cidade Universitária, Ilha do Fundão, Rio de Janeiro - RJ – Brazil. E-mail: [email protected] 2Laboratório de Ecologia de Insetos, Department of Ecologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, A0-113, Bloco A, Av. Carlos Chagas Filho, 373, Cidade Universitária ,Ilha do Fundão, Rio de Janeiro - RJ – Brazil Abstract Here we describe a new Photinina genus with three species endemic to Serra dos Órgãos Mountains in Brazil. Ybytyra- moan gen. nov. occurs in high altitudes, from 980m up to 2000m, and has the following unique set of characters: head abruptly depressed at vertex; lanterns not fully developed, somewhat rounded or anteriorly rounded, straight posteriad, with posterolateral rounded projections (billycock-shaped), at the middle of the abdominal sterna VI and VII; abdominal sternum VIII not covered by VII; phallus and parameres apically teethed. We provide illustrations and a key to the three species in this genus: Ybytyramoan praeclarum sp. nov. (type-species), Y. diasi sp. -
Tve429 Archangelsky.Qxp
MIGUEL ARCHANGELSKY Universidad Nacional de La Patagonia, CONICET, Esquel, Chubut, Argentina DESCRIPTION OF THE LAST LARVAL INSTAR AND PUPA OF ASPISOMA FENESTRATA BLANCHARD, 1837 (COLEOPTERA: LAMPYRIDAE) WITH BRIEF NOTES ON ITS BIOLOGY Archangelsky, M., 2004. Description of the last larval instar and pupa of Aspisoma fenestrata Blanchard, 1839 (Coleoptera: Lampyridae) with brief notes on its biology. – Tijdschrift voor Entomologie 147: 49-56, figs. 1-8, tables 1-2. [ISSN 0040-7496]. Published 1 June 2004. The last instar larva and pupa of Aspisoma fenestrata are described and figured for the first time. Notes for comparison with two other unidentified Aspisoma larvae are provided, as well as brief notes on the biology of A. fenestrata. Comparison of Aspisoma larvae with other known Crato- morphini larvae places Aspisoma closer to Pyractomena than to Cratomorphus. Correspondence: M. Archangelsky, Laboratorio de Ecología Acuática (LEA-CONICET); Univer- sidad Nacional de La Patagonia; Sarmiento 849; 9200 Esquel, Chubut; Argentina. E-mail: hy- [email protected]. Key words. – Lampyridae; fireflies; Aspisoma; larvae; Neotropical. There are over 40 genera of fireflies in the Neotrop- MATERIAL AND METHODS ical region, most of which are present in South Amer- ica. Surprisingly, this contrasts with the very few de- Two larvae were collected from inside a rotting log scriptions of South American lampyrid larvae and partially immersed in a pool of saline temporary water pupae. Up to now the only published descriptions are gathered at the sides of a dirt road connecting the lo- those by Costa et al. (1988) and Viviani (1989). In cality of Totoralejos with Rd. 60. This locality is with- their book, Costa et al. -
Firefly Official Insect of Pennsylvania
Official insect of the Commonwealth of Pennsylvania Firefly Photuris pennsylvanica Adopted: April 10, 1974 Firefly: Official insect of the Commonwealth of Pennsylvania This file is licensed under the Wikipedia Creative Commons Attribution 2.0 Generic license. Pennsylvania Law The following information was excerpted from the The Pennsylvania Statutes, Title 71, Chapter 6, Section 1010. Title 71 P.S. State Government I. The Administrative Codes and Related Provisions Chapter 6. Provisions Similar or Closely Related to Provisions of the Administrative Code Secretary and Department of Internal Affairs State Emblems § 1010. State insect The firefly (Lampyridae Coleoptera) of the species Photuris pensylvanica De Geer is hereby selected, designated and adopted as the official insect of the Commonwealth of Pennsylvania. CREDIT(S) 1974, April 10, P.L. 247, No. 59, § 1. As amended 1988, Dec. 5, P.L. 1101, No. 130, § 1, effective in 60 days. HISTORICAL AND STATUTORY NOTES 1990 Main Volume The 1988 amendment substituted "(Lampyridae Coleoptera)" of the species Photuris pensylvanica De Geer" for "(Lampyridae)". Title of Act: An Act selecting, designating and adopting the firefly as the official insect of the Commonwealth of Pennsylvania. 1974, April 10, P.L. 247, No. 59. 71 P.S. § 1010, PA ST 71 P.S. § 1010 Sources... Thomson Reuters: Westlaw The Pennsylvania Statutes, <http://government.westlaw.com/linkedslice/default.asp?SP=pac-1000> (Accessed August 10, 2010) Shearer, Benjamin F. and Barbara S. State Names, Seals, Flags and Symbols: A Historical Guide Third Edition, Revised and Expanded. Westport, Conn: Greenwood Press, 3 Sub edition, 2001. Pennsylvania State Insect Firefly (Photinus Pyralsis) Adopted on April 10, 1974. -
Ent17 4 367 402 (Kazantsev Perez-Gelabert).Pmd
Russian Entomol. J. 17(4): 367402 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2008 Fireflies of Hispaniola (Coleoptera: Lampyridae) Ñâåòëÿ÷êè îñòðîâà Ãàèòè (Coleoptera: Lampyridae) Sergey V. Kazantsev1 & Daniel E. Perez-Gelabert2 Ñ.Â. Êàçàíöåâ1, Ä.E. Ïåðåñ-Ãåëàáåðò2 1 Insect Centre, Donetskaya 13326, Moscow 109651, Russia 1 Èíñåêò-öåíòð, óë. Äîíåöêàÿ 13326, Ìîñêâà 109651, Ðîññèÿ, E-mail: [email protected] 2 Department of Entomology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC 205637012, USA, E-mail: [email protected] KEY WORDS: Coleoptera, Lampyridae, new species, taxonomy, Greater Antilles, Neotropics. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Coleoptera, Lampyridae, íîâûå âèäû, òàêñîíîìèÿ, Áîëüøèå Àíòèëüñêèå îñòðîâà, Íåîòðîïèêà. ABSTRACT. Thirty three new fireflies, Lychnacris mirabilis Kazantsev et Perez-Gelabert, 2009 spp.n. â atrocrocea, L. bahorucoensis, L. cienagaensis, L. hier- îñíîâíîì èç êîëëåêöèé Èíñòèòóòà áîòàíè÷åñêèõ è roi, L. montensis, L. orbis, L. piceonotata, L. rufocaer- çîîëîãè÷åñêèõ èññëåäîâàíèé ïðè Óíèâåðñèòåòå Ñàí- ulea, L. scintilla, Callopisma altimontana, C. domini- òî-Äîìèíãî è Íàöèîíàëüíîãî ìóçåÿ åñòåñòâåííîé cana, C. engombe, C. lamellicornis, C. larimarena, C. èñòîðèè Ñàíòî Äîìèíãî. Ðîäà Callopisma Motschul- rubicunda, Erythrolychnia azuensis, E. caborojensis, sky, 1853 è Erythrolychnia Motschulsky, 1853 ïåðåíî- E. cristobalensis, E. marcanoi, E. medranoi, E. peder- ñÿòñÿ èç òðèáû Photinini â òðèáó Cratomorphini. Ery- nalensis, E. roseimargo, Robopus acutangulus, R. bas- throlychnia olivieri Leng et Mutchler, 1922 syn.n. tardoi, R. dissimilis, R. hondovallensis, R. nigrifrons, ñâîäèòñÿ â ñèíîíèìû ê E. bipartita (E. Olivier, 1912). R. peregrinus, R. vallinovae, Heterophotinus montico- Ïðèâîäèòñÿ ïîëíûé ñïèñîê ëàìïèðèä îñòðîâà Ãàèòè la, H. nubilus, H. striatus and Presbyolampis mirabilis âìåñòå ñ êàðòàìè àðåàëîâ, à òàêæå îïðåäåëèòåëüíûå Kazantsev et Perez-Gelabert, 2009 spp.n., are described òàáëèöû äëÿ òðèá, ðîäîâ è âèäîâ. -
Redalyc.Bioluminescent Coleoptera of Biological Station of Boracéia
Biota Neotropica ISSN: 1676-0611 [email protected] Instituto Virtual da Biodiversidade Brasil Viviani, Vadim Ravara; Machado dos Santos, Raphael Bioluminescent Coleoptera of Biological Station of Boracéia (Salesópolis, SP, Brazil): diversity, bioluminescence and habitat distribution Biota Neotropica, vol. 12, núm. 3, septiembre, 2012, pp. 1-14 Instituto Virtual da Biodiversidade Campinas, Brasil Available in: http://www.redalyc.org/articulo.oa?id=199124391001 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Biota Neotrop., vol. 12, no. 3 Bioluminescent Coleoptera of Biological Station of Boracéia (Salesópolis, SP, Brazil): diversity, bioluminescence and habitat distribution Vadim Ravara Viviani1,2 & Raphael Machado dos Santos1 1Laboratório de Bioquímica e Biotecnologia de Sistemas Bioluminescentes, Graduate School of Biotechnology and Environmental Monitoring, Universidade Federal de São Carlos – UFSCar, Campus de Sorocaba, Rod. João Leme dos Santos, Km 110, Itinga, CEP 18052-780, Sorocaba, SP, Brazil 2Corresponding author: Vadim Ravara Viviani, e-mail: [email protected] Viviani, V.R. & Santos R.M. Bioluminescent Coleoptera of Biological Station of Boracéia (Salesópolis, SP, Brazil): diversity, bioluminescence and habitat distribution. Biota Neotrop. 12(3): http://www.biotaneotropica. org.br/v12n3/en/abstract?article+bn00212032012 Abstract: Brazil hosts the richest biodiversity of bioluminescent beetles in the world. Several species are found in the Atlantic rain forest, one of the richest and most threatened tropical forests in the world. We have catalogued the biodiversity of bioluminescent species mainly of Elateroidea superfamily occurring in one of the last largest and most preserved remnants of Atlantic rain forest, located at the Biological Station of Boracéia of São Paulo University (Salesopolis, SP, Brazil). -
Coleoptera: Lampyridae) with New Ohio Records and Regional Observations for Several Firefly Species
Ohio Biological Survey Notes 9: 16–34, 2019. © Ohio Biological Survey, Inc. Life History and Updated Range Extension of Photinus scintillans (Coleoptera: Lampyridae) with New Ohio Records and Regional Observations for Several Firefly Species LYNN F. FAUST¹, LAURA S. HUGHES2, MARK H. ZLOBA3, AND HEATHER L. FARRINGTON4 1Lynn F. Faust, 11828 Couch Mill Rd, Knoxville, TN 37932, [email protected]; 2Laura S. Hughes, 365 Shawnee Loop South, Pataskala, Ohio. [email protected]; 3Mark H. Zloba, Ecological Manager, Cincinnati Museum Center, Edge of Appalachia Preserve System, 4274 Waggoner Riffle Rd., West Union, Ohio 45693, [email protected]; 4Heather L. Farrington, Cincinnati Museum Center, Curator of Zoology, 1301 Western Avenue, Cincinnati, Ohio 45203, [email protected]. Abstract: Photinus scintillans (Say) has long been considered the Photinus species with one of the smallest ranges in North America. In field studies conducted between 2016 and 2019 in Ohio and Indiana, we discovered new, thriving P. scintillans populations, tripling the east-west range from 550 km to 1820 km when combined with more recent collection records by firefly researchers Lloyd, Stanger-Hall, and Lower. We describe in new detail flight behaviors, nocturnal timing of activity, flash pattern, lantern coloration changes, courtship, and mating habits. We present the first evidence of the presence of spermatophore-producing spiral glands and prolonged mating with the brachypterous females; oviposition behaviors; larval eclosion and appearance; and seasonality with habitat variations and commonalities. We provide the first report with photos of possible phoresy by a springtail (Collembola) on a firefly. In addition, we offer new Ohio state (and nearby Indiana and Kentucky) firefly records, including the extremely rare P. -
A Molecular Phylogeny of Lampyridae with Insight Into Visual and Bioluminescent Evolution
Brigham Young University BYU ScholarsArchive Theses and Dissertations 2014-12-01 A Molecular Phylogeny of Lampyridae with Insight into Visual and Bioluminescent Evolution Gavin Jon Martin Brigham Young University - Provo Follow this and additional works at: https://scholarsarchive.byu.edu/etd Part of the Biology Commons BYU ScholarsArchive Citation Martin, Gavin Jon, "A Molecular Phylogeny of Lampyridae with Insight into Visual and Bioluminescent Evolution" (2014). Theses and Dissertations. 5758. https://scholarsarchive.byu.edu/etd/5758 This Thesis is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. A Molecular Phylogeny of Lampyridae with Insight into Visual and Bioluminescent Evolution Gavin J. Martin A thesis submitted to the faculty of Brigham Young University in partial fulfillment of the requirements for the degree of Master of Science Seth M. Bybee, Chair Michael F. Whiting Marc A. Branham Department of Biology Brigham Young University December 2014 Copyright © 2014 Gavin J. Martin All Rights Reserved ABSTRACT A Molecular Phylogeny of Lampyridae with Insight into Visual and Bioluminescent Evolution Gavin J. Martin Department of Biology, BYU Master of Science Fireflies are some of the most captivating organisms on the planet. Because of this, they have a rich history of study, especially concerning their bioluminescent and visual behavior. Among insects, opsin copy number variation has been shown to be quite diverse. However, within the beetles, very little work on opsins has been conducted. Here we look at the visual system of fireflies (Coleoptera: Lampyridae), which offer an elegant system in which to study visual evolution as it relates to their behavior and broader ecology. -
Bioluminescent Coleoptera of Biological Station of Boracéia (Salesópolis, SP, Brazil): Diversity, Bioluminescence and Habitat Distribution
Bioluminescent Coleoptera of Biological Station of Boracéia (Salesópolis, SP, Brazil): diversity, bioluminescence and habitat distribution Viviani, V.R. & Santos R.M. Biota Neotrop. 2012, 12(3): 000-000. On line version of this paper is available from: http://www.biotaneotropica.org.br/v12n3/en/abstract?article+bn00212032012 A versão on-line completa deste artigo está disponível em: http://www.biotaneotropica.org.br/v12n3/pt/abstract?article+bn00212032012 Received/ Recebido em 16/09/11 - Revised/ Versão reformulada recebida em 26/04/12 - Accepted/ Publicado em 02/07/12 ISSN 1676-0603 (on-line) Biota Neotropica is an electronic, peer-reviewed journal edited by the Program BIOTA/FAPESP: The Virtual Institute of Biodiversity. This journal’s aim is to disseminate the results of original research work, associated or not to the program, concerned with characterization, conservation and sustainable use of biodiversity within the Neotropical region. Biota Neotropica é uma revista do Programa BIOTA/FAPESP - O Instituto Virtual da Biodiversidade, que publica resultados de pesquisa original, vinculada ou não ao programa, que abordem a temática caracterização, conservação e uso sustentável da biodiversidade na região Neotropical. Biota Neotropica is an eletronic journal which is available free at the following site http://www.biotaneotropica.org.br A Biota Neotropica é uma revista eletrônica e está integral e gratuitamente disponível no endereço http://www.biotaneotropica.org.br Biota Neotrop., vol. 12, no. 3 Bioluminescent Coleoptera of Biological Station of Boracéia (Salesópolis, SP, Brazil): diversity, bioluminescence and habitat distribution Vadim Ravara Viviani1,2 & Raphael Machado dos Santos1 1Laboratório de Bioquímica e Biotecnologia de Sistemas Bioluminescentes, Graduate School of Biotechnology and Environmental Monitoring, Universidade Federal de São Carlos – UFSCar, Campus de Sorocaba, Rod. -
Information to Users
INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough. substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author (fid not send UMI a complete manuscript and there are missing pages, these wilt be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Photographs included in the original manuscript have been reproduced xerographicaity in this copy. Higher quality 6’ x 9“ black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. ProQuest Information and Learning 300 North Zeeb Road. Ann Arbor, Ml 48106-1346 USA 800-521-0600 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. THE EVOLUTION OF LAMPYRIDAE, WITH SPECIAL EMPHASIS ON THE ORIGIN OF PHOTIC BEHAVIOR AND SIGNAL SYSTEM EVOLUTION (COLEOPTERA: LAMPYRIDAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for The Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Marc A. -
A Summary of the Endemic Beetle Genera of the West Indies (Insecta: Coleoptera); Bioindicators of the Evolutionary Richness of This Neotropical Archipelago
University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Center for Systematic Entomology, Gainesville, Insecta Mundi Florida 2-29-2012 A summary of the endemic beetle genera of the West Indies (Insecta: Coleoptera); bioindicators of the evolutionary richness of this Neotropical archipelago Stewart B. Peck Carleton University, [email protected] Daniel E. Perez-Gelabert Department of Entomology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC 20013- 7012. USA, [email protected] Follow this and additional works at: https://digitalcommons.unl.edu/insectamundi Part of the Entomology Commons Peck, Stewart B. and Perez-Gelabert, Daniel E., "A summary of the endemic beetle genera of the West Indies (Insecta: Coleoptera); bioindicators of the evolutionary richness of this Neotropical archipelago" (2012). Insecta Mundi. 718. https://digitalcommons.unl.edu/insectamundi/718 This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. INSECTA MUNDI A Journal of World Insect Systematics 0212 A summary of the endemic beetle genera of the West Indies (Insecta: Coleoptera); bioindicators of the evolutionary richness of this Neotropical archipelago Stewart B. Peck Department of Biology Carleton University 1125 Colonel By Drive Ottawa, ON K1S 5B6, Canada Daniel E. Perez-Gelabert Department of Entomology U. S. National Museum of Natural History, Smithsonian Institution P. O. Box 37012 Washington, D. C., 20013-7012, USA Date of Issue: February 29, 2012 CENTER FOR SYSTEMATIC ENTOMOLOGY, INC., Gainesville, FL Stewart B. -
A Day-Flashing Photinus Firefly (Coleoptera: Lampyridae) from Central Panamá: an Emergent Shift to Predator-Free Space?
Insect Systematics & Evolution (2017) DOI 10.1163/1876312X-48022162 brill.com/ise A day-flashing Photinus firefly (Coleoptera: Lampyridae) from central Panamá: an emergent shift to predator-free space? Fredric V. Vencla,b,*, Xin Luanc, Xinhua Fuc and Luana S. Marojad aDepartment of Ecology and Evolution, Stony Brook University, Stony Brook, NY 11794–5245, USA bNational Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA cCollege of Plant Science & Technology, Huazhong Agricultural University, Wuhan 430070, P.R. China dDepartment of Biology, Williams College, 31 Morley Drive, Williamstown, MA 01267, USA *Corresponding author, e-mail: [email protected] Abstract Fireflies in the genus Photinus are well regarded for their luminescent nocturnal courtship displays. Here we report on a new firefly species,Photinus interdius, which is remarkable for its fully diurnal and lu- minescent courtship protocol. Males slowly flew near the ground searching for receptive females and emitted 800 ms, bright yellow light flashes at 3–4-s intervals. Male flights occurred as early as 13:10 and ceased before 18:00. We sequenced two mitochondrial loci and one genomic locus and combined these with those from 99 specimens representing 45 Photinus and 25 related firefly species. Bayesian inference resulted in a well-resolved phylogeny that placed this new species as the closest relative of, but basal to the Photinus clade. We propose that the adaptive significance of this extraordinary temporal shift in courtship niche is the outcome of a selective landscape that has optimized the trade-off between reduced predation risk and ease of mate-localization. Keywords aedeagus; Bayesian; bioluminescence; courtship; photometry; phylogeny; predation; sexual selection Introduction Fireflies of the exclusively New World genus Photinus are well-known for their spec- tacular nocturnal courtship dialogues, wherein flying males broadcast bright flashes of light to locate conspecific, sedentary females, who emit flashed responses with species- specific time delays.