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| authority = [[Constantine Samuel Rafinesque|Rafinesque]], 1815 | | authority = [[Constantine Samuel Rafinesque|Rafinesque]], 1815 | ||
| subdivision_ranks = Subfamilies | | subdivision_ranks = Subfamilies | ||
| subdivision = Amydetinae<ref name=Martin2019>{{cite journal |last1=Martin |first1=Gavin J. |last2=Stanger-Hall |first2=Kathrin F. |last3=Branham | first3=Marc A. |last4=Da Silveira |first4=Luiz F. L. |last5=Lower |first5=Sarah E. |last6=Hall |first6=David W. |last7=Li |first7=Xue-Yan |last8=Lemmon |first8=Alan R. |last9=Moriarty Lemmon |first9=Emily |last10=Bybee |first10=Seth M. |display-authors=3 |editor-last=Jordal |editor-first=Bjarte |title=Higher-Level Phylogeny and Reclassification of Lampyridae (Coleoptera: Elateroidea) |journal=[[Insect Systematics and Diversity]] |publisher=[[Oxford University Press]] ) |volume=3 |issue=6 |date=1 November 2019 |doi=10.1093/isd/ixz024 }}</ref><br/>[[Cheguevariinae]]<ref name=Ferreira2019>{{cite journal |last1=Ferreira |first1=Vinicius S. |last2=Keller |first2=Oliver |last3=Branham |first3=Marc A. |last4=Ivie |first4=Michael A. |title=Molecular data support the placement of the enigmatic ''Cheguevaria'' as a subfamily of Lampyridae (Insecta: Coleoptera) |journal=[[Zoological Journal of the Linnean Society]] |volume=187 |issue=4 |pages=1253–1258 |publisher=[[Oxford University Press]] |year=2019 |doi=10.1093/zoolinnean/zlz073 |doi-access=free }}</ref><br/>[[Chespiritoinae]]<ref name="FerreiraEtAl">{{cite journal |last1=Ferreira |first1=Vinicius S. |last2=Keller |first2=Oliver |last3=Branham |first3=Marc A |editor-last=Marvaldi |editor-first=Adriana |title=Multilocus Phylogeny Support the Nonbioluminescent Firefly Chespirito as a New Subfamily in the Lampyridae (Coleoptera: Elateroidea) |journal=[[Insect Systematics and Diversity]] |publisher=[[Oxford University Press]] |volume=4 |issue=6 |date=1 November 2020 |doi=10.1093/isd/ixaa014 }}</ref><br/> | | subdivision = Amydetinae<ref name=Martin2019>{{cite journal |last1=Martin |first1=Gavin J. |last2=Stanger-Hall |first2=Kathrin F. |last3=Branham | first3=Marc A. |last4=Da Silveira |first4=Luiz F. L. |last5=Lower |first5=Sarah E. |last6=Hall |first6=David W. |last7=Li |first7=Xue-Yan |last8=Lemmon |first8=Alan R. |last9=Moriarty Lemmon |first9=Emily |last10=Bybee |first10=Seth M. |display-authors=3 |editor-last=Jordal |editor-first=Bjarte |title=Higher-Level Phylogeny and Reclassification of Lampyridae (Coleoptera: Elateroidea) |journal=[[Insect Systematics and Diversity]] |publisher=[[Oxford University Press]] ) |volume=3 |issue=6 |date=1 November 2019 |article-number=11 |doi=10.1093/isd/ixz024 }}</ref><br/>[[Cheguevariinae]]<ref name=Ferreira2019>{{cite journal |last1=Ferreira |first1=Vinicius S. |last2=Keller |first2=Oliver |last3=Branham |first3=Marc A. |last4=Ivie |first4=Michael A. |title=Molecular data support the placement of the enigmatic ''Cheguevaria'' as a subfamily of Lampyridae (Insecta: Coleoptera) |journal=[[Zoological Journal of the Linnean Society]] |volume=187 |issue=4 |pages=1253–1258 |publisher=[[Oxford University Press]] |year=2019 |doi=10.1093/zoolinnean/zlz073 |doi-access=free }}</ref><br/>[[Chespiritoinae]]<ref name="FerreiraEtAl">{{cite journal |last1=Ferreira |first1=Vinicius S. |last2=Keller |first2=Oliver |last3=Branham |first3=Marc A |editor-last=Marvaldi |editor-first=Adriana |title=Multilocus Phylogeny Support the Nonbioluminescent Firefly Chespirito as a New Subfamily in the Lampyridae (Coleoptera: Elateroidea) |journal=[[Insect Systematics and Diversity]] |publisher=[[Oxford University Press]] |volume=4 |issue=6 |date=1 November 2020 |article-number=2 |doi=10.1093/isd/ixaa014 }}</ref><br/> | ||
[[Cyphonocerinae]]<br/> | [[Cyphonocerinae]]<br/> | ||
Lamprohizinae<ref name=Martin2019/><br/> | Lamprohizinae<ref name=Martin2019/><br/> | ||
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}} | }} | ||
The '''Lampyridae''' are a [[family (biology)|family]] of [[Elateroidea|elateroid]] beetles with more than 2, | The '''Lampyridae''' are a [[family (biology)|family]] of [[Elateroidea|elateroid]] beetles with more than 2,400 described species,<ref name="COL-BRN">{{Cite web |title=Catalogue of Life: taxon Lampyridae |url=https://www.catalogueoflife.org/data/taxon/BRN |website=Catalogue of Life |language=en |access-date=2025-09-05}}</ref> many of which are [[bioluminescence|light-emitting]]. They are soft-bodied beetles commonly called '''fireflies''', '''lightning bugs''', or '''[[glowworm]]s''' for their conspicuous production of light, mainly [[crepuscular|during twilight]], to attract mates.<ref>[https://bugguide.net/node/view/85#names BugGuide: common names]</ref> The [[type species]] is ''[[Lampyris noctiluca]]'', the common glow-worm of Europe. Light production in the Lampyridae is thought to have originated as a [[Aposematism|warning]] [[honest signal|signal]] that the [[larva]]e were distasteful. This ability to create light was then co-opted as a mating signal and, in a further development, adult female fireflies of the genus ''[[Photuris]]'' mimic the flash pattern of the ''[[Photinus (beetle)|Photinus]]'' beetle to trap their males as prey. | ||
Fireflies are found in [[temperate]] and [[tropical]] climates. Many live in [[marsh]]es or in wet, wooded areas where their larvae have abundant sources of food. Although all | Fireflies are found in [[temperate]] and [[tropical]] climates. Many live in [[marsh]]es or in wet, wooded areas where their larvae have abundant sources of food. Although all fireflies nominally glow as larvae, only some species produce light in their adult stage, and the location of the light organ varies among species and between sexes of the same species. Fireflies have attracted human attention since [[classical antiquity]]; their presence has been taken to signify a wide variety of conditions in different cultures and is especially appreciated aesthetically in Japan, where parks are set aside for this specific purpose.<!--this is a summary of cited claims in the main text--> | ||
==Biology== | ==Biology== | ||
[[File:Luciola4 crop.png|thumb|left|A [[larviform female]] with light-emitting organs on her [[abdomen]]. Unlike actual larvae, she has [[compound eye]]s.]] | [[File:Luciola4 crop.png|thumb|left|A [[larviform female]] with light-emitting organs on her [[abdomen]]. Unlike actual larvae, she has [[compound eye]]s.]] | ||
Fireflies are | Fireflies are beetles and in many aspects resemble other beetles at all stages of their [[biological life cycle|life cycle]], undergoing [[complete metamorphosis]].<ref name="Lewis 2016"/> A few days after mating, a female lays her fertilized eggs on or just below the surface of the ground. The eggs hatch three to four weeks later.<ref name="Marshall 2020"/> In certain firefly species with aquatic larvae, such as ''[[Aquatica leii]]'', the female oviposits on emergent portions of aquatic plants, and the larvae descend into the water after hatching.<ref>{{cite journal |last1=Fu |first1=Xinhua |last2=Nobuyoshi |first2=Ohba |last3=Vencl |first3=Fredric V. |last4=Lei |first4=Chaoliang |title=Life cycle and behaviour of the aquatic firefly ''Luciola leii'' (Coleoptera: Lampyridae) from Mainland China |journal=The Canadian Entomologist |year=2006 |volume=138 |issue=6 |pages=860–870 |doi=10.4039/n05-093 |s2cid=84624340 |url=https://www.researchgate.net/publication/250370510}}</ref> | ||
The larvae feed until the end of the summer. Most fireflies [[insect winter ecology|hibernate]] as larvae. Some do this by burrowing underground, while others find places on or under the bark of trees. They emerge in the spring. At least one species, ''[[Ellychnia corrusca]]'', overwinters as an adult.<ref>{{cite web |url=https://extension.umaine.edu/home-and-garden-ipm/fact-sheets/common-name-listing/winter-firefly/ |title=Winter Firefly |website=The University of Maine Cooperative Extension |last=Armstrong |first=Charles |access-date=22 June 2022}}</ref> The larvae of most species are specialized [[predator]]s and feed on other larvae, terrestrial [[snail]]s, and [[slug]]s. Some are so specialized that they have grooved [[mandible (insect)|mandibles]] that deliver digestive fluids directly to their prey. The larval stage lasts from several weeks up to, in certain species, two or more years.<ref>{{cite web |url=https://www.xerces.org/endangered-species/fireflies/about |title=About Fireflies |website=Xerces Society for Invertebrate Conservation |date=17 November 2021 |access-date=22 June 2022}}</ref> The larvae [[pupa]]te for one to two and a half weeks and emerge as adults.<ref name="Marshall 2020"/> | The larvae feed until the end of the summer. Most fireflies [[insect winter ecology|hibernate]] as larvae. Some do this by burrowing underground, while others find places on or under the bark of trees. They emerge in the spring. At least one species, ''[[Ellychnia corrusca]]'', overwinters as an adult.<ref>{{cite web |url=https://extension.umaine.edu/home-and-garden-ipm/fact-sheets/common-name-listing/winter-firefly/ |title=Winter Firefly |website=The University of Maine Cooperative Extension |last=Armstrong |first=Charles |access-date=22 June 2022}}</ref> The larvae of most species are specialized [[predator]]s and feed on other larvae, terrestrial [[snail]]s, and [[slug]]s. Some are so specialized that they have grooved [[mandible (insect)|mandibles]] that deliver digestive fluids directly to their prey. The larval stage lasts from several weeks up to, in certain species, two or more years.<ref>{{cite web |url=https://www.xerces.org/endangered-species/fireflies/about |title=About Fireflies |website=Xerces Society for Invertebrate Conservation |date=17 November 2021 |access-date=22 June 2022}}</ref> The larvae [[pupa]]te for one to two and a half weeks and emerge as adults.<ref name="Marshall 2020"/> | ||
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Adult diet varies among firefly species: some are predatory, while others feed on plant [[pollen]] or [[nectar]]. Some adults, like the European glow-worm, have no mouth, emerging only to mate and lay eggs before dying. In most species, adults live for a few weeks in summer.<ref name="Marshall 2020">{{cite news |last=Marshall |first=Michael |title=Why the lights are going out for fireflies |url=https://www.theguardian.com/environment/2020/feb/22/why-lights-going-out-fireflies-conservation-pollution |access-date=4 February 2022 |work=[[The Guardian]] |date=22 February 2020}}</ref><ref name="McLean Buck Hanson 1972">{{cite journal |last1=McLean |first1=Miriam |last2=Buck |first2=John |last3=Hanson |first3=Frank E. |title=Culture and Larval Behavior of Photurid Fireflies |journal=[[The American Midland Naturalist]] |volume=87 |issue=1 |publisher=[[University of Notre Dame]] |year=1972 |pages=133–145 |doi=10.2307/2423887 |jstor=2423887 }}</ref> | Adult diet varies among firefly species: some are predatory, while others feed on plant [[pollen]] or [[nectar]]. Some adults, like the European glow-worm, have no mouth, emerging only to mate and lay eggs before dying. In most species, adults live for a few weeks in summer.<ref name="Marshall 2020">{{cite news |last=Marshall |first=Michael |title=Why the lights are going out for fireflies |url=https://www.theguardian.com/environment/2020/feb/22/why-lights-going-out-fireflies-conservation-pollution |access-date=4 February 2022 |work=[[The Guardian]] |date=22 February 2020}}</ref><ref name="McLean Buck Hanson 1972">{{cite journal |last1=McLean |first1=Miriam |last2=Buck |first2=John |last3=Hanson |first3=Frank E. |title=Culture and Larval Behavior of Photurid Fireflies |journal=[[The American Midland Naturalist]] |volume=87 |issue=1 |publisher=[[University of Notre Dame]] |year=1972 |pages=133–145 |doi=10.2307/2423887 |jstor=2423887 }}</ref> | ||
Fireflies vary widely in their general appearance, with differences in color, shape, size, and features such as antennae. Adults differ in size depending on the species, with the largest up to {{convert|1|in|mm|0|order=flip|abbr=on}} long. Many species have non-flying [[larviform female]]s. These can often be distinguished from the larvae only because the adult females have [[arthropod eye|compound eyes]], unlike the [[Simple eye in invertebrates|simple eye]]s of larvae, though the females have much smaller (and often highly regressed) eyes than those of their males.<ref name="Lau & Meyer-Rochow 2006">{{cite journal |last1=Lau |first1=T. F. |last2=Meyer-Rochow |first2=V. B. |date=2006 |title=Sexual dimorphism in the compound eye of Rhagophthalmus ohbai (Coleoptera: Rhagophthalmidae): Morphology and ultrastructure |journal=Journal of Asia-Pacific Entomology |volume=9 |issue=1 |pages=19–30 |doi=10.1016/S1226-8615(08)60271-X|bibcode=2006JAsPE...9...19L }}</ref> The most commonly known fireflies are [[Nocturnality|nocturnal]], although numerous species are [[diurnality|diurnal]] and usually not luminescent; however, some species that remain in shadowy areas may produce light.<ref name="Lewis 2016">{{cite book |last=Lewis |first=Sara |title=Silent Sparks: The Wondrous World of Fireflies |isbn=978- | Fireflies vary widely in their general appearance, with differences in color, shape, size, and features such as antennae. Adults differ in size depending on the species, with the largest up to {{convert|1|in|mm|0|order=flip|abbr=on}} long. Many species have non-flying [[larviform female]]s. These can often be distinguished from the larvae only because the adult females have [[arthropod eye|compound eyes]], unlike the [[Simple eye in invertebrates|simple eye]]s of larvae, though the females have much smaller (and often highly regressed) eyes than those of their males.<ref name="Lau & Meyer-Rochow 2006">{{cite journal |last1=Lau |first1=T. F. |last2=Meyer-Rochow |first2=V. B. |date=2006 |title=Sexual dimorphism in the compound eye of Rhagophthalmus ohbai (Coleoptera: Rhagophthalmidae): Morphology and ultrastructure |journal=Journal of Asia-Pacific Entomology |volume=9 |issue=1 |pages=19–30 |doi=10.1016/S1226-8615(08)60271-X|bibcode=2006JAsPE...9...19L }}</ref> The most commonly known fireflies are [[Nocturnality|nocturnal]], although numerous species are [[diurnality|diurnal]] and usually not luminescent; however, some species that remain in shadowy areas may produce light.<ref name="Lewis 2016">{{cite book |last=Lewis |first=Sara |title=Silent Sparks: The Wondrous World of Fireflies |isbn=978-1-4008-8031-7 |publisher=Princeton University Press |url=https://books.google.com/books?id=i9-BCwAAQBAJ&q=Firefly+metamorphosis&pg=PA17 |date=26 April 2016 |page=17}}</ref> | ||
Most fireflies are distasteful to vertebrate predators, as they contain the steroid pyrones [[lucibufagin]]s, similar to the [[cardiotonic]] [[Bufanolide|bufadienolides]] found in some poisonous toads.<ref>{{Cite journal |last1=Eisner |first1=Thomas |last2=Wiemer |first2=David |last3=Haynes |first3=Leroy |last4=Meinwald |first4=Jerrold |title=Lucibufagins: Defensive steroids from the fireflies Photinus ignitus and P. marginellus (Coleoptera: Lampyridae) |pmid=16592501 |year=1978 |volume=75 |issue=2 |pages=905–908 |pmc=411366 |journal=[[PNAS]] |doi=10.1073/pnas.75.2.905 |bibcode=1978PNAS...75..905E |doi-access=free }}</ref> All fireflies glow as larvae, where bioluminescence is an [[Aposematism|aposematic]] warning signal to [[predator]]s.<ref name="Lewis Cratsley 2008">{{cite journal |last1=Lewis |first1=Sara M. |last2=Cratsley |first2=Christopher K. |s2cid=16360536 |date=January 2008 |title=Flash Signal Evolution, Mate Choice, and Predation in Fireflies |journal=[[Annual Review of Entomology]] |volume=53 |issue=1 |pages=293–321 |doi=10.1146/annurev.ento.53.103106.093346 |pmid=17877452}}</ref><ref>{{Cite journal |last1=Branham |first1=Marc A. |last2=Wenzel |first2=John W. |date=December 2001 |title=The Evolution of Bioluminescence in Cantharoids (Coleoptera: Elateroidea) |journal=[[The Florida Entomologist]] |volume=84 |issue=4 | | Most fireflies are distasteful to vertebrate predators, as they contain the steroid pyrones [[lucibufagin]]s, similar to the [[cardiotonic]] [[Bufanolide|bufadienolides]] found in some poisonous toads.<ref>{{Cite journal |last1=Eisner |first1=Thomas |last2=Wiemer |first2=David |last3=Haynes |first3=Leroy |last4=Meinwald |first4=Jerrold |title=Lucibufagins: Defensive steroids from the fireflies Photinus ignitus and P. marginellus (Coleoptera: Lampyridae) |pmid=16592501 |year=1978 |volume=75 |issue=2 |pages=905–908 |pmc=411366 |journal=[[PNAS]] |doi=10.1073/pnas.75.2.905 |bibcode=1978PNAS...75..905E |doi-access=free }}</ref> All fireflies glow as larvae, where bioluminescence is an [[Aposematism|aposematic]] warning signal to [[predator]]s.<ref name="Lewis Cratsley 2008">{{cite journal |last1=Lewis |first1=Sara M. |last2=Cratsley |first2=Christopher K. |s2cid=16360536 |date=January 2008 |title=Flash Signal Evolution, Mate Choice, and Predation in Fireflies |journal=[[Annual Review of Entomology]] |volume=53 |issue=1 |pages=293–321 |doi=10.1146/annurev.ento.53.103106.093346 |pmid=17877452}}</ref><ref>{{Cite journal |last1=Branham |first1=Marc A. |last2=Wenzel |first2=John W. |date=December 2001 |title=The Evolution of Bioluminescence in Cantharoids (Coleoptera: Elateroidea) |journal=[[The Florida Entomologist]] |volume=84 |issue=4 |page=565 |doi=10.2307/3496389 |jstor=3496389 |url=http://journals.fcla.edu/flaent/article/view/75005 |doi-access=free|url-access=subscription }}</ref>{{sfn|Gullan|Cranston|2014|pp=121–122}} | ||
=== Light and chemical production === | === Light and chemical production === | ||
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[[File:Firefly composite.jpg|thumb|left|''[[Photuris]]'' female by flash photography (above); by her own light (below)]] | [[File:Firefly composite.jpg|thumb|left|''[[Photuris]]'' female by flash photography (above); by her own light (below)]] | ||
Light production in fireflies is due to the chemical process of [[bioluminescence]]. This occurs in specialized light-emitting [[organ (biology)|organ]]s, usually on a female firefly's lower [[Abdomen (insect anatomy)|abdomen]]. | Light production in fireflies is due to the chemical process of [[bioluminescence]]. This occurs in specialized light-emitting [[organ (biology)|organ]]s (known as [[photophore]]s), usually located on a female firefly's lower [[Abdomen (insect anatomy)|abdomen]]. These fireflies produce the enzyme [[firefly luciferase]] that acts on another compound produced by fireflies, [[firefly luciferin]]. In the presence of [[magnesium]] ions, [[Adenosine triphosphate|ATP]], and oxygen, this reaction produces light.<ref name="pmid8805542">{{cite journal |vauthors=Baldwin TO |date=March 1996 |title=Firefly luciferase: the structure is known, but the mystery remains |journal=Structure |volume=4 |issue=3 |pages=223–228 |doi=10.1016/S0969-2126(96)00026-3 |pmid=8805542 |doi-access=free}}</ref><ref name="oba">{{cite journal |vauthors=Oba Y, Ojika M, Inouye S |date=April 2003 |title=Firefly luciferase is a bifunctional enzyme: ATP-dependent monooxygenase and a long chain fatty acyl-CoA synthetase |journal=FEBS Letters |volume=540 |issue=1–3 |pages=251–254 |bibcode=2003FEBSL.540..251O |doi=10.1016/S0014-5793(03)00272-2 |pmid=12681517 |s2cid=12075190 |doi-access=free}}</ref> Oxygen is supplied via an abdominal trachea or breathing tube. [[Gene]] coding for these substances has been [[Luciferase#Applications|inserted into many different organisms]].<ref name="Day 2009">{{cite book |last=Day |first=John |chapter=Beetle bioluminescence: a genetic and enzymatic research review |year=2009 |publisher=Research Signpost: Kerala |editor=Meyer-Rochow, V. B. |title=Bioluminescence in Focus |pages=325–355 }}</ref> Firefly luciferase is used in [[forensics]], and the enzyme has medical uses – in particular, for detecting the presence of [[Adenosine triphosphate|ATP]] or magnesium.<ref name="Lewis Cratsley 2008"/> Fireflies produce a "cold light", with no [[infrared]] or [[ultraviolet]] frequencies. The light may be yellow, green, or pale red, with [[wavelength]]s from 510 to 670 nanometers. Some species such as the dimly glowing "[[Phausis reticulata|blue ghost]]" of the Eastern US may seem to emit blueish-white light from a distance and in low light conditions, but their glow is bright green when observed up close.<ref name="Frick-Ruppert_Rosen_2008">{{Cite journal |last1=Frick-Ruppert |first1=Jennifer E. |last2=Rosen |first2=Joshua J. |date=2008 |title=Morphology and Behavior of Phausis Reticulata (Blue Ghost Firefly) |journal=Journal of the North Carolina Academy of Science |volume=124 |issue=4 |pages=139–147}}</ref> Their perceived blue tint may be due to the [[Purkinje effect]].<ref>{{cite journal |last1=Branchini |first1=Bruce R. |last2=Southworth |first2=Tara L. |last3=Salituro |first3=Leah J. |last4=Fontaine |first4=Danielle M. |last5=Oba |first5=Yuichi |title=Cloning of the Blue Ghost (Phausis reticulata) Luciferase Reveals a Glowing Source of Green Light |journal=[[Photochemistry and Photobiology]] |date=2017 |volume=93 |issue=2 |pages=473–478 |pmid=27696431 |doi=10.1111/php.12649 |doi-access=free }}</ref> During a study on the genome of ''Aquatica leii'', scientists discovered two key genes are responsible for the formation, activation, and positioning of this firefly's light organ: Alabd-B and AlUnc-4.<ref>{{Cite journal |last1=Fu |first1=Xinhua |last2=Zhu |first2=Xinlei |date=2024-03-05 |title=Key homeobox transcription factors regulate the development of the firefly's adult light organ and bioluminescence |journal=Nature Communications |language=en |volume=15 |issue=1 |page=1736 |doi=10.1038/s41467-024-45559-7 |pmid=38443352 |issn=2041-1723|pmc=10914744 |bibcode=2024NatCo..15.1736F }}</ref> | ||
Adults emit light primarily for mate selection. Early larval bioluminescence was adopted in the phylogeny of adult fireflies, and was repeatedly gained and lost before becoming fixed and retained as a mechanism of sexual communication in many species.<ref name="Lewis Cratsley 2008"/><ref>{{cite journal |last1=Martin |first1=Gavin J. |last2=Branham |first2=Marc A. |last3=Whiting |first3=Michael F. |last4=Bybee |first4=Seth M. |date=February 2017 |title=Total evidence phylogeny and the evolution of adult bioluminescence in fireflies (Coleoptera: Lampyridae) |journal=[[Molecular Phylogenetics and Evolution]] |volume=107 |pages=564–575 |doi=10.1016/j.ympev.2016.12.017 |pmid=27998815 |doi-access=free|bibcode=2017MolPE.107..564M }}</ref> Adult lampyrids have a variety of ways to communicate with mates in courtships: steady glows, flashing, and the use of chemical signals unrelated to photic systems.<ref>{{cite journal |last1=Stanger-Hall |first1=K. F. |last2=Lloyd |first2=J. E. |last3=Hillis |first3=D. M. |title=Phylogeny of North American fireflies (Coleoptera: Lampyridae): implications for the evolution of light signals |journal=[[Molecular Phylogenetics and Evolution]] |volume=45 |issue=1 |pages=33–49 |doi=10.1016/j.ympev.2007.05.013 |year=2007 |pmid=17644427|bibcode=2007MolPE..45...33S }}</ref> Chemical signals, or pheromones, are the ancestral form of sexual communication; this pre-dates the evolution of flash signaling in the lineage, and is retained today in diurnally-active species.<ref name="Lewis Cratsley 2008" /><ref>{{cite journal |last=Branham |first=M. |date=February 2003 |title=The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae) |journal=[[Cladistics (journal)|Cladistics]] |volume=19 |issue=1 |pages=1–22 |doi=10.1111/j.1096-0031.2003.tb00404.x |pmid=34905865 |s2cid=46266960 }}</ref> Some species, especially lightning bugs of the [[genera]] ''[[Photinus (beetle)|Photinus]]'', ''[[Photuris (genus)|Photuris]]'', and ''[[Pyractomena]]'', are distinguished by the unique courtship flash patterns emitted by flying males in search of females. In general, females of the genus ''Photinus'' do not fly, but do give a flash response to males of their own species. Signals, whether photic or chemical, allow fireflies to identify mates of their own species. Flash signaling characteristics include differences in duration, timing, color, number and rate of repetitions, height of flight, and direction of flight (e.g. climbing or diving) and vary interspecifically and geographically.<ref name=":2">{{Cite journal |last1=Stanger-Hall |first1=Kathrin F. |last2=Lloyd |first2=James E. |date=March 2015 |title=Flash signal evolution in Photinus fireflies: Character displacement and signal exploitation in a visual communication system |journal=[[Evolution (journal)|Evolution]] |volume=69 |issue=3 |pages=666–682 |doi=10.1111/evo.12606 |pmid=25627920 |s2cid=26075485 |doi-access=free }}</ref>{{sfn|Gullan|Cranston|2014|pp=121–122}} When flash signals are not sufficiently distinguished between species in a population, [[sexual selection]] encourages divergence of signaling patterns.<ref name=":2"/> | Adults emit light primarily for mate selection. Early larval bioluminescence was adopted in the phylogeny of adult fireflies, and was repeatedly gained and lost before becoming fixed and retained as a mechanism of sexual communication in many species.<ref name="Lewis Cratsley 2008"/><ref>{{cite journal |last1=Martin |first1=Gavin J. |last2=Branham |first2=Marc A. |last3=Whiting |first3=Michael F. |last4=Bybee |first4=Seth M. |date=February 2017 |title=Total evidence phylogeny and the evolution of adult bioluminescence in fireflies (Coleoptera: Lampyridae) |journal=[[Molecular Phylogenetics and Evolution]] |volume=107 |pages=564–575 |doi=10.1016/j.ympev.2016.12.017 |pmid=27998815 |doi-access=free|bibcode=2017MolPE.107..564M }}</ref> Adult lampyrids have a variety of ways to communicate with mates in courtships: steady glows, flashing, and the use of chemical signals unrelated to photic systems.<ref>{{cite journal |last1=Stanger-Hall |first1=K. F. |last2=Lloyd |first2=J. E. |last3=Hillis |first3=D. M. |title=Phylogeny of North American fireflies (Coleoptera: Lampyridae): implications for the evolution of light signals |journal=[[Molecular Phylogenetics and Evolution]] |volume=45 |issue=1 |pages=33–49 |doi=10.1016/j.ympev.2007.05.013 |year=2007 |pmid=17644427|bibcode=2007MolPE..45...33S }}</ref> Chemical signals, or pheromones, are the ancestral form of sexual communication; this pre-dates the evolution of flash signaling in the lineage, and is retained today in diurnally-active species.<ref name="Lewis Cratsley 2008" /><ref>{{cite journal |last=Branham |first=M. |date=February 2003 |title=The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae) |journal=[[Cladistics (journal)|Cladistics]] |volume=19 |issue=1 |pages=1–22 |doi=10.1111/j.1096-0031.2003.tb00404.x |pmid=34905865 |s2cid=46266960 }}</ref> Some species, especially lightning bugs of the [[genera]] ''[[Photinus (beetle)|Photinus]]'', ''[[Photuris (genus)|Photuris]]'', and ''[[Pyractomena]]'', are distinguished by the unique courtship flash patterns emitted by flying males in search of females. In general, females of the genus ''Photinus'' do not fly, but do give a flash response to males of their own species. Signals, whether photic or chemical, allow fireflies to identify mates of their own species. Flash signaling characteristics include differences in duration, timing, color, number and rate of repetitions, height of flight, and direction of flight (e.g. climbing or diving) and vary interspecifically and geographically.<ref name=":2">{{Cite journal |last1=Stanger-Hall |first1=Kathrin F. |last2=Lloyd |first2=James E. |date=March 2015 |title=Flash signal evolution in Photinus fireflies: Character displacement and signal exploitation in a visual communication system |journal=[[Evolution (journal)|Evolution]] |volume=69 |issue=3 |pages=666–682 |doi=10.1111/evo.12606 |pmid=25627920 |bibcode=2015Evolu..69..666S |s2cid=26075485 |doi-access=free }}</ref>{{sfn|Gullan|Cranston|2014|pp=121–122}} When flash signals are not sufficiently distinguished between species in a population, [[sexual selection]] encourages divergence of signaling patterns.<ref name=":2"/> | ||
Synchronization of flashing occurs in several species; it is explained as [[phase synchronization]] and spontaneous order.<ref>{{cite book |last=Murray |first=James D. |url=https://books.google.com/books?id=1QM3h80gb_IC |title=Mathematical Biology |publisher=[[Springer Science+Business Media|Springer]] |year=2002 |isbn=978-0-387-95223-9 |edition=3rd |volume=I. An Introduction |pages=295–299 |language=en |author-link=James D. Murray}}</ref> Tropical fireflies routinely synchronise their flashes among large groups, particularly in Southeast Asia. At night along river banks in the Malaysian jungles, fireflies synchronize their light emissions precisely. | Synchronization of flashing occurs in several species; it is explained as [[phase synchronization]] and spontaneous order.<ref>{{cite book |last=Murray |first=James D. |url=https://books.google.com/books?id=1QM3h80gb_IC |title=Mathematical Biology |publisher=[[Springer Science+Business Media|Springer]] |year=2002 |isbn=978-0-387-95223-9 |edition=3rd |volume=I. An Introduction |pages=295–299 |language=en |author-link=James D. Murray}}</ref> Tropical fireflies routinely synchronise their flashes among large groups, particularly in Southeast Asia. At night along river banks in the Malaysian jungles, fireflies synchronize their light emissions precisely. Hypotheses for the causes of this behavior involve diet, social interaction, and altitude. In the Philippines, thousands of fireflies can be seen all year-round in the town of [[Donsol]]. In the United States, one of the most famous sightings of fireflies blinking in [[unison]] occurs annually near [[Elkmont, Tennessee]], in the [[Great Smoky Mountains]] during the first weeks of June.<ref>[https://www.nps.gov/grsm/learn/nature/fireflies.htm Synchronous Fireflies – Great Smoky Mountains National Park]. Nps.gov (3 June 2013). Retrieved on 22 June 2013.</ref> [[Congaree National Park]] in [[South Carolina]] is another host to this phenomenon.<ref>Cross, Robert (23 May 2004) "[https://web.archive.org/web/20050318055636/http://www.seacoastonline.com/2004news/05232004/travel/17745.htm Tree huggin']". ''Chicago Tribune''.</ref> | ||
Female "femme fatale" ''Photuris'' fireflies mimic the photic signaling patterns of the smaller ''Photinus'', attracting males to what appears to be a suitable mate, then eating them.<ref name="Lewis Cratsley 2008"/> This provides the females with a supply of the toxic defensive lucibufagin chemicals.{{sfn|Gullan|Cranston|2014|p=387}} | Female "femme fatale" ''Photuris'' fireflies mimic the photic signaling patterns of the smaller ''Photinus'', attracting males to what appears to be a suitable mate, then eating them.<ref name="Lewis Cratsley 2008"/> This provides the females with a supply of the toxic defensive lucibufagin chemicals.{{sfn|Gullan|Cranston|2014|p=387}} | ||
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=== Fossil history === | === Fossil history === | ||
The oldest known fossils of the Lampyridae family are ''Protoluciola'' and ''Flammarionella'' from the [[Late Cretaceous]] ([[Cenomanian]] ~ 99 million years ago) [[Burmese amber]] of Myanmar, which belong to the subfamily [[Luciolinae]]. The light producing organs are clearly present.<ref name="Kazantsev 2015">{{Cite journal |last=Kazantsev |first=S. V. |date=December 2015 |title=Protoluciola albertalleni gen.n., sp.n., a new Luciolinae firefly (Insecta: Coleoptera: Lampyridae) from Burmite amber |url=http://kmkjournals.com/journals/REJ/REJ_Index_Volumes/REJ_24/REJ_24_4_281_283_Kazantsev |journal=Russian Entomological Journal |volume=24 |issue=1 |pages=281–283 |doi=10.15298/rusentj.24.4.02 |doi-access=free}}</ref><ref>{{Cite journal |last1=Cai |first1=Chenyang |last2=Tihelka |first2=Erik |last3=Ballantyne |first3=Lesley |last4=Li |first4=Yan-Da |last5=Huang |first5=Diying |last6=Engel |first6=Michael S. |last7=Kundrata |first7=Robin |date=January 2024 |title=A light in the dark: a mid-Cretaceous bioluminescent firefly with specialized antennal sensory organs |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=291 |issue=2030 |doi=10.1098/rspb.2024.1671 |issn=0962-8452 |pmc=11387053 |pmid=39255838 | The oldest known fossils of the Lampyridae family are ''Protoluciola'' and ''Flammarionella'' from the [[Late Cretaceous]] ([[Cenomanian]] ~ 99 million years ago) [[Burmese amber]] of Myanmar, which belong to the subfamily [[Luciolinae]]. The light producing organs are clearly present.<ref name="Kazantsev 2015">{{Cite journal |last=Kazantsev |first=S. V. |date=December 2015 |title=Protoluciola albertalleni gen.n., sp.n., a new Luciolinae firefly (Insecta: Coleoptera: Lampyridae) from Burmite amber |url=http://kmkjournals.com/journals/REJ/REJ_Index_Volumes/REJ_24/REJ_24_4_281_283_Kazantsev |journal=Russian Entomological Journal |volume=24 |issue=1 |pages=281–283 |doi=10.15298/rusentj.24.4.02 |doi-access=free}}</ref><ref>{{Cite journal |last1=Cai |first1=Chenyang |last2=Tihelka |first2=Erik |last3=Ballantyne |first3=Lesley |last4=Li |first4=Yan-Da |last5=Huang |first5=Diying |last6=Engel |first6=Michael S. |last7=Kundrata |first7=Robin |date=January 2024 |title=A light in the dark: a mid-Cretaceous bioluminescent firefly with specialized antennal sensory organs |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=291 |issue=2030 |article-number=20241671 |doi=10.1098/rspb.2024.1671 |issn=0962-8452 |pmc=11387053 |pmid=39255838}}</ref> The ancestral glow colour for the last common ancestor of all living fireflies has been inferred to be green, based on [[genomics|genomic analysis]].<ref name="Oba 2020">{{Cite journal |last1=Oba |first1=Y. |last2=Konishi |first2=K. |last3=Yano |first3=D. |last4=Shibata |first4=H. |last5=Kato |first5=D. |last6=Shirai |first6=T. |date=December 2020 |title=Resurrecting the ancient glow of the fireflies |journal=[[Science Advances]] |volume=6 |issue=49 |article-number=eabc5705 |doi=10.1126/sciadv.abc5705 |pmc=7710365 |pmid=33268373 |bibcode=2020SciA....6.5705O}}</ref> | ||
=== Taxonomy === | === Taxonomy === | ||
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}} | }} | ||
|2={{clade | |2={{clade | ||
|1=Lamprohizinae [[File:Lamprohiza splendidula01 (cropped).jpg|80px]] | |1=[[Lamprohizinae]] [[File:Lamprohiza splendidula01 (cropped).jpg|80px]] | ||
|2={{clade | |2={{clade | ||
|1={{clade | |1={{clade | ||
| Line 141: | Line 140: | ||
[[File:Fireflies, Georgia, US.jpg|thumb|center|upright=3.6|Fireflies in [[Georgia (U.S. state)|Georgia (USA)]], 8-second exposure]] | [[File:Fireflies, Georgia, US.jpg|thumb|center|upright=3.6|Fireflies in [[Georgia (U.S. state)|Georgia (USA)]], 8-second exposure]] | ||
Firefly populations are thought to be declining worldwide. While monitoring data for many regions are scarce, a growing number of anecdotal reports, coupled with several published studies from Europe and Asia, suggest that fireflies are endangered.<ref>{{Cite journal |last1=Atkins |first1=Val |last2=Bell |first2=Dolly |last3=Bowker |first3=Ann |last4=Charig |first4=Marcus |last5=Crew |first5=June |last6=Dale |first6=Martin |last7=Hickmott |first7=Bill |last8=Payne |first8=Barbara |last9=Pendleton |first9=Dilys |last10=Pendleton |first10=Trevor |last11=Robinson |first11=Martin |display-authors=3 |date=2016 |title=The status of the glow-worm Lampyris noctiluca L. (Coleoptera: Lampyridae) in England |url=https://www.researchgate.net/publication/333000808 |journal=Lampyrid |volume=4 |pages=20–35}}</ref><ref>{{Cite journal |last1=Gardiner |first1=Tim |last2=Didham |first2=Raphael K. |date=2020 |title=Glowing, glowing, gone? Monitoring long-term trends in glow-worm numbers in south-east England |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/icad.12407 |journal=Insect Conservation and Diversity |language=en |volume=13 |issue=2 |pages=162–174 |doi=10.1111/icad.12407 |s2cid=216387774 |url-access=subscription }}</ref><ref>{{Cite journal |last1=Khoo |first1=Veronica |last2=Nada |first2=B. |last3=Kirton |first3=L. G. |last4=Chooi-Khim |first4=Phon |display-authors=3 |date=2009 |title=Monitoring the population of the firefly Pteroptyx tener along the Selangor River, Malaysia for conservation and sustainable ecotourism |url=https://www.researchgate.net/publication/285742573 |journal=Lampyrid |volume=2 |pages=162–173}}</ref><ref>{{Cite journal |last1=Wong |first1=C. |last2=Yeap |first2=C. A. |date=2012 |title=Conservation of congregating firefly zones (CFZs) in peninsular Malaysia |journal=Lampyrid |volume=2 |pages=174–187}}</ref> Recent [[IUCN Red List]] assessments for North American fireflies have identified species with heightened extinction risk in the US, with 18 taxa categorized as threatened with extinction.<ref>{{Cite web |title=Latest Update to the IUCN Red List Includes First Global Assessments for Fireflies, with a Spotlight on North America {{!}} Xerces Society |url=https://xerces.org/blog/iucn-red-list-assess-fireflies |access-date=2021-10-12 |website=xerces.org}}</ref><ref>{{Cite journal |last1=Fallon |first1=Candace E. |last2=Walker |first2=Anna C. |last3=Lewis |first3=Sara |last4=Cicero |first4=Joseph |last5=Faust |first5=Lynn |last6=Heckscher |first6=Christopher M. |last7=Pérez-Hernández |first7=Cisteil X. |last8=Pfeiffer |first8=Ben |last9=Jepsen |first9=Sarina |display-authors=3 |date=2021-11-17 |title=Evaluating firefly extinction risk: Initial red list assessments for North America |journal=[[PLOS One]] |volume=16 |issue=11 | | Firefly populations are thought to be declining worldwide. While monitoring data for many regions are scarce, a growing number of anecdotal reports, coupled with several published studies from Europe and Asia, suggest that fireflies are endangered.<ref>{{Cite journal |last1=Atkins |first1=Val |last2=Bell |first2=Dolly |last3=Bowker |first3=Ann |last4=Charig |first4=Marcus |last5=Crew |first5=June |last6=Dale |first6=Martin |last7=Hickmott |first7=Bill |last8=Payne |first8=Barbara |last9=Pendleton |first9=Dilys |last10=Pendleton |first10=Trevor |last11=Robinson |first11=Martin |display-authors=3 |date=2016 |title=The status of the glow-worm Lampyris noctiluca L. (Coleoptera: Lampyridae) in England |url=https://www.researchgate.net/publication/333000808 |journal=Lampyrid |volume=4 |pages=20–35}}</ref><ref>{{Cite journal |last1=Gardiner |first1=Tim |last2=Didham |first2=Raphael K. |date=2020 |title=Glowing, glowing, gone? Monitoring long-term trends in glow-worm numbers in south-east England |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/icad.12407 |journal=Insect Conservation and Diversity |language=en |volume=13 |issue=2 |pages=162–174 |doi=10.1111/icad.12407 |s2cid=216387774 |url-access=subscription }}</ref><ref>{{Cite journal |last1=Khoo |first1=Veronica |last2=Nada |first2=B. |last3=Kirton |first3=L. G. |last4=Chooi-Khim |first4=Phon |display-authors=3 |date=2009 |title=Monitoring the population of the firefly Pteroptyx tener along the Selangor River, Malaysia for conservation and sustainable ecotourism |url=https://www.researchgate.net/publication/285742573 |journal=Lampyrid |volume=2 |pages=162–173}}</ref><ref>{{Cite journal |last1=Wong |first1=C. |last2=Yeap |first2=C. A. |date=2012 |title=Conservation of congregating firefly zones (CFZs) in peninsular Malaysia |journal=Lampyrid |volume=2 |pages=174–187}}</ref> Recent [[IUCN Red List]] assessments for North American fireflies have identified species with heightened extinction risk in the US, with 18 taxa categorized as threatened with extinction.<ref>{{Cite web |title=Latest Update to the IUCN Red List Includes First Global Assessments for Fireflies, with a Spotlight on North America {{!}} Xerces Society |url=https://xerces.org/blog/iucn-red-list-assess-fireflies |access-date=2021-10-12 |website=xerces.org}}</ref><ref>{{Cite journal |last1=Fallon |first1=Candace E. |last2=Walker |first2=Anna C. |last3=Lewis |first3=Sara |last4=Cicero |first4=Joseph |last5=Faust |first5=Lynn |last6=Heckscher |first6=Christopher M. |last7=Pérez-Hernández |first7=Cisteil X. |last8=Pfeiffer |first8=Ben |last9=Jepsen |first9=Sarina |display-authors=3 |date=2021-11-17 |title=Evaluating firefly extinction risk: Initial red list assessments for North America |journal=[[PLOS One]] |volume=16 |issue=11 |article-number=e0259379 |doi=10.1371/journal.pone.0259379 |pmid=34788329 |pmc=8598072 |bibcode=2021PLoSO..1659379F |doi-access=free }}</ref> | ||
Fireflies face threats including habitat loss and degradation, [[light pollution]], [[pesticide]] use, poor water quality, invasive species, over-collection, and [[climate change]].<ref>{{Cite journal |last1=Lewis |first1=Sara M. |last2=Wong |first2=Choong Hay |last3=Owens |first3=Avalon C. S. |last4=Fallon |first4=Candace |last5=Jepsen |first5=Sarina |last6=Thancharoen |first6=Anchana |last7=Wu |first7=Chiahsiung |last8=De Cock |first8=Raphael |last9=Novák |first9=Martin |last10=López-Palafox |first10=Tania |last11=Khoo |first11=Veronica |display-authors=3 |date=1 February 2020 |title=A Global Perspective on Firefly Extinction Threats |journal=[[BioScience]] |volume=70 |issue=2 |pages=157–167 |doi=10.1093/biosci/biz157 |doi-access=free}}</ref> Firefly tourism, a quickly growing sector of the travel and tourism industry, has also been identified as a potential threat to fireflies and their habitats when not managed appropriately.<ref>{{Cite journal |last1=Lewis |first1=Sara M. |last2=Thancharoen |first2=Anchana |last3=Wong |first3=Choong Hay |last4=López-Palafox |first4=Tania |last5=Santos |first5=Paola Velasco |last6=Wu |first6=Chiahsiung |last7=Faust |first7=Lynn |last8=Cock |first8=Raphaël De |last9=Owens |first9=Avalon C. S. |last10=Lemelin |first10=R. Harvey |last11=Gurung |first11=Hum |display-authors=3 |date=2021 |title=Firefly tourism: Advancing a global phenomenon toward a brighter future |journal=Conservation Science and Practice |volume=3 |issue=5 |article-number=e391 |doi=10.1111/csp2.391 |doi-access=free|bibcode=2021ConSP...3E.391L |hdl=10067/1773950151162165141 |hdl-access=free }}</ref> Like many other organisms, fireflies are directly affected by land-use change (e.g., loss of habitat area and connectivity), which is identified as the main driver of [[biodiversity]] changes in terrestrial ecosystems.<ref>{{Cite journal |last1=Sala |first1=Osvaldo E. |last2=Chapin |first2=F. Stuart III |last3=Armesto |first3=Juan J. |last4=Berlow |first4=Eric |last5=Bloomfield |first5=Janine |last6=Dirzo |first6=Rodolfo |last7=Huber-Sanwald |first7=Elisabeth |last8=Huenneke |first8=Laura F. |display-authors=3 |date=2000-03-10 |title=Global Biodiversity Scenarios for the Year 2100 |journal=[[Science (journal)|Science]] |volume=287 |issue=5459 |pages=1770–1774 |doi=10.1126/science.287.5459.1770 |pmid=10710299|bibcode=2000Sci...287.1770S |s2cid=13336469 }}</ref> Pesticides, including [[insecticide]]s and [[herbicide]]s, have also been indicated as a likely cause of firefly decline.<ref>Potter, Daniel A. See [https://www.firefly.org/how-you-can-help.html "How You Can Help"], FireFly.org, citing (1) "Understanding Halofenozide (Mach 2) and Imidacloprid (Merit) Soil Insecticides," International SportsTurf Institute, Incorporated, Turfax, Vol. 6, No. 1 (Jan-Feb 1998) and (2) "Relative Toxicities of Chemicals to the Earthworm Eisenia foetida," by Brian L. Roberts and H. Wyman Dorough. Article first published online: 20 October 2009. Environmental Toxicology and Chemistry, Vol. 3, No. 1 (Jan. 1984), pp. 67–78.</ref><ref name=":3">{{Cite web |last1=Fallon |first1=Candace |last2=Hoyle |first2=Sarah |last3=Lewis |first3=Sara |last4=Owens |first4=Avalon |last5=Lee-Mader |first5=Eric |last6=Hoffman Black |first6=Scott |last7=Jepsen |first7=Sarina |display-authors=3 |year=2019 |title=Conserving the Jewels of the Night: Guidelines for Protecting Fireflies in the United States and Canada |url=https://xerces.org/sites/default/files/publications/19-049_01_Firefly%20guidelines_web.pdf |access-date=2021-06-23 |website=The Xerces Society for Invertebrate Conservation |location=Portland, Oregon |page=}}</ref> These chemicals can not only harm fireflies directly but also potentially reduce prey populations and degrade habitat. Light pollution is an especially concerning threat to fireflies. Since the majority of firefly species use bioluminescent courtship signals,<ref name="Lloyd 1989 373">{{Cite journal |last1=Lloyd |first1=James E. |last2=Wing |first2=Steven R. |last3=Hongtrakul |first3=Tawatchai |title=Ecology, Flashes, and Behavior of Congregating Thai Fireflies |journal=[[Biotropica]] |volume=21 |issue=4 |doi=10.2307/2388290 |page=373 |jstor=2388290 |year=1989|bibcode=1989Biotr..21..373L }}</ref> they are also sensitive to environmental levels of light and consequently to [[light pollution]].<ref name="Lloyd 1989 373"/><ref name=":0">{{Cite journal |last1=Viviani |first1=Vadim Ravara |last2=Rocha |first2=Mayra Yamazaki |last3=Hagen |first3=Oskar |date=June 2010 |title=Fauna de besouros bioluminescentes (Coleoptera: Elateroidea: Lampyridae; Phengodidae, Elateridae) nos municípios de Campinas, Sorocaba-Votorantim e Rio Claro-Limeira (SP, Brasil): biodiversidade e influência da urbanização |journal=Biota Neotropica |volume=10 |issue=2 |pages=103–116 |doi=10.1590/s1676-06032010000200013 |doi-access=free}}</ref> A growing number of studies investigating the effects of artificial light at night on fireflies has shown that light pollution can disrupt fireflies' courtship signals and even interfere with larval dispersal.<ref>{{Cite journal |last1=Firebaugh |first1=Ariel |last2=Haynes |first2=Kyle J. |date=2016-12-01 |title=Experimental tests of light-pollution impacts on nocturnal insect courtship and dispersal |journal=[[Oecologia]] |volume=182 |issue=4 |pages=1203–1211 |doi=10.1007/s00442-016-3723-1 |pmid=27646716 |bibcode=2016Oecol.182.1203F |s2cid=36670391}}</ref><ref>{{Cite journal |last1=Owens |first1=Avalon Celeste Stevahn |last2=Meyer-Rochow |first2=Victor Benno |last3=Yang |first3=En-Cheng |date=7 February 2018 |title=Short- and mid-wavelength artificial light influences the flash signals of Aquatica ficta fireflies (Coleoptera: Lampyridae) |journal=PLOS ONE |volume=13 |issue=2 |article-number=e0191576 |doi=10.1371/journal.pone.0191576 |pmid=29415023 |pmc=5802884 |bibcode=2018PLoSO..1391576O |doi-access=free}}</ref><ref>{{Cite journal |last1=Owens |first1=Avalon C. S. |last2=Lewis |first2=Sara M. |date=2021 |title=Narrow-spectrum artificial light silences female fireflies (Coleoptera: Lampyridae) |url=https://onlinelibrary.wiley.com/doi/10.1111/icad.12487 |journal=Insect Conservation and Diversity |volume=14 |issue=2 |pages=199–210 |doi=10.1111/icad.12487 |s2cid=232246018 |url-access=subscription }}</ref><ref>{{Cite journal |last1=Owens |first1=Avalon C. S. |last2=Lewis |first2=Sara M. |date=2021 |title=Effects of artificial light on growth, development, and dispersal of two North American fireflies (Coleoptera: Lampyridae) |url=https://linkinghub.elsevier.com/retrieve/pii/S002219102100010X |journal=Journal of Insect Physiology |volume=130 |article-number=104200 |doi=10.1016/j.jinsphys.2021.104200 |pmid=33607160 |bibcode=2021JInsP.13004200O |s2cid=231969942|url-access=subscription }}</ref> Researchers agree that protecting and enhancing firefly habitat is necessary to conserve their populations. Recommendations include reducing or limiting artificial light at night, restoring habitats where threatened species occur, and eliminating unnecessary pesticide use, among many others.<ref name=":3"/><ref>{{Cite web |title=How You Can Help Prevent Fireflies from Disappearing |url=https://www.firefly.org/how-you-can-help.html |access-date=12 October 2021 |website=Firefly.org }}</ref><ref>{{Cite web |title=How You Can Help |publisher=[[Xerces Society]] |url=https://xerces.org/endangered-species/fireflies/how-you-can-help |access-date=12 October 2021}}</ref> | |||
=== In culture === | === In culture === | ||
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In Italy, the firefly ([[Italian language|Italian]]: {{lang|it|lucciola}}) appears in Canto XXVI of [[Inferno (Dante)|Dante's ''Inferno'']], written in the 14th century:<ref>{{cite book |last=Alighieri |first=Dante |author-link=Dante Alighieri |title=[[Inferno (Dante)|Inferno]] |date=1320 |at=Canto XXVI, lines 25–32}}</ref> | In Italy, the firefly ([[Italian language|Italian]]: {{lang|it|lucciola}}) appears in Canto XXVI of [[Inferno (Dante)|Dante's ''Inferno'']], written in the 14th century:<ref>{{cite book |last=Alighieri |first=Dante |author-link=Dante Alighieri |title=[[Inferno (Dante)|Inferno]] |date=1320 |at=Canto XXVI, lines 25–32}}</ref> | ||
{{blockquote|<poem>Quante | {{blockquote|<poem>Quante 'l villan ch'al poggio si riposa, | ||
nel tempo che colui che | nel tempo che colui che 'l mondo schiara | ||
la faccia sua a noi tien meno ascosa, | la faccia sua a noi tien meno ascosa, | ||
come la mosca cede a la zanzara, | come la mosca cede a la zanzara, | ||
vede lucciole giù per la vallea, | vede lucciole giù per la vallea, | ||
forse colà | forse colà dov' e' vendemmia e ara: | ||
di tante fiamme tutta risplendea | di tante fiamme tutta risplendea | ||
l'ottava bolgia, ...</poem>|Dante's ''Inferno'', Canto XXVI, lines 25–32}} | |||
{{blockquote|As many as the fireflies which the peasant sees in the <nowiki>[</nowiki>[[Tuscany|Tuscan]]<nowiki>]</nowiki> valley below, when he is resting on the hill—in the season [midsummer] when the sun hides least from us, and at the time of day [dusk] when the fly gives place to the mosquito—perhaps in the fields where he tills the ground and gathers in the grapes; with that many flames the eighth ditch [of Hell] was shining, ...|prose translation}} | {{blockquote|As many as the fireflies which the peasant sees in the <nowiki>[</nowiki>[[Tuscany|Tuscan]]<nowiki>]</nowiki> valley below, when he is resting on the hill—in the season [midsummer] when the sun hides least from us, and at the time of day [dusk] when the fly gives place to the mosquito—perhaps in the fields where he tills the ground and gathers in the grapes; with that many flames the eighth ditch [of Hell] was shining, ...|prose translation}} | ||
| Line 184: | Line 181: | ||
* Faust, Lynn Frierson (2017). [https://ugapress.org/book/9780820348728/fireflies-glow-worms-and-lightning-bugs/ "Fireflies, Glow-worms, and Lightning Bugs"] | * Faust, Lynn Frierson (2017). [https://ugapress.org/book/9780820348728/fireflies-glow-worms-and-lightning-bugs/ "Fireflies, Glow-worms, and Lightning Bugs"] | ||
* {{cite journal |last1=Lewis |first1=S. M. |first2=C. K. |last2=Cratsley |s2cid=16360536 |year=2008 |title=Flash signal evolution, mate choice, and predation in fireflies |journal=[[Annual Review of Entomology]] |volume=53 |pages=293–321 |doi=10.1146/annurev.ento.53.103106.093346 |pmid=17877452 |ref=none }} | * {{cite journal |last1=Lewis |first1=S. M. |first2=C. K. |last2=Cratsley |s2cid=16360536 |year=2008 |title=Flash signal evolution, mate choice, and predation in fireflies |journal=[[Annual Review of Entomology]] |volume=53 |pages=293–321 |doi=10.1146/annurev.ento.53.103106.093346 |pmid=17877452 |ref=none }} | ||
* [[Sara Lewis|Lewis, Sara]] (2016). Silent Sparks: The Wondrous World of Fireflies. Princeton University Press. | * [[Sara Lewis|Lewis, Sara]] (2016). Silent Sparks: The Wondrous World of Fireflies. Princeton University Press. {{ISBN|978-1-4008-8031-7}} | ||
* {{cite web |last=Stous |first=Hollend |year=1997 |title=A review of predation in ''Photuris'', and its effects on the evolution of flash signaling in other New World fireflies |url=http://www.colostate.edu/Depts/Entomology/courses/en507/papers_1997/stous.html |ref=none }} | * {{cite web |last=Stous |first=Hollend |year=1997 |title=A review of predation in ''Photuris'', and its effects on the evolution of flash signaling in other New World fireflies |url=http://www.colostate.edu/Depts/Entomology/courses/en507/papers_1997/stous.html |ref=none }} | ||
Latest revision as of 08:29, 16 November 2025
Template:Short description Script error: No such module "about". Script error: No such module "redirect hatnote". Template:Good article Template:Use dmy dates Template:Automatic taxobox
The Lampyridae are a family of elateroid beetles with more than 2,400 described species,[1] many of which are light-emitting. They are soft-bodied beetles commonly called fireflies, lightning bugs, or glowworms for their conspicuous production of light, mainly during twilight, to attract mates.[2] The type species is Lampyris noctiluca, the common glow-worm of Europe. Light production in the Lampyridae is thought to have originated as a warning signal that the larvae were distasteful. This ability to create light was then co-opted as a mating signal and, in a further development, adult female fireflies of the genus Photuris mimic the flash pattern of the Photinus beetle to trap their males as prey.
Fireflies are found in temperate and tropical climates. Many live in marshes or in wet, wooded areas where their larvae have abundant sources of food. Although all fireflies nominally glow as larvae, only some species produce light in their adult stage, and the location of the light organ varies among species and between sexes of the same species. Fireflies have attracted human attention since classical antiquity; their presence has been taken to signify a wide variety of conditions in different cultures and is especially appreciated aesthetically in Japan, where parks are set aside for this specific purpose.
Biology
Fireflies are beetles and in many aspects resemble other beetles at all stages of their life cycle, undergoing complete metamorphosis.[3] A few days after mating, a female lays her fertilized eggs on or just below the surface of the ground. The eggs hatch three to four weeks later.[4] In certain firefly species with aquatic larvae, such as Aquatica leii, the female oviposits on emergent portions of aquatic plants, and the larvae descend into the water after hatching.[5]
The larvae feed until the end of the summer. Most fireflies hibernate as larvae. Some do this by burrowing underground, while others find places on or under the bark of trees. They emerge in the spring. At least one species, Ellychnia corrusca, overwinters as an adult.[6] The larvae of most species are specialized predators and feed on other larvae, terrestrial snails, and slugs. Some are so specialized that they have grooved mandibles that deliver digestive fluids directly to their prey. The larval stage lasts from several weeks up to, in certain species, two or more years.[7] The larvae pupate for one to two and a half weeks and emerge as adults.[4]
Adult diet varies among firefly species: some are predatory, while others feed on plant pollen or nectar. Some adults, like the European glow-worm, have no mouth, emerging only to mate and lay eggs before dying. In most species, adults live for a few weeks in summer.[4][8]
Fireflies vary widely in their general appearance, with differences in color, shape, size, and features such as antennae. Adults differ in size depending on the species, with the largest up to Template:Convert long. Many species have non-flying larviform females. These can often be distinguished from the larvae only because the adult females have compound eyes, unlike the simple eyes of larvae, though the females have much smaller (and often highly regressed) eyes than those of their males.[9] The most commonly known fireflies are nocturnal, although numerous species are diurnal and usually not luminescent; however, some species that remain in shadowy areas may produce light.[3]
Most fireflies are distasteful to vertebrate predators, as they contain the steroid pyrones lucibufagins, similar to the cardiotonic bufadienolides found in some poisonous toads.[10] All fireflies glow as larvae, where bioluminescence is an aposematic warning signal to predators.[11][12]Template:Sfn
Light and chemical production
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Light production in fireflies is due to the chemical process of bioluminescence. This occurs in specialized light-emitting organs (known as photophores), usually located on a female firefly's lower abdomen. These fireflies produce the enzyme firefly luciferase that acts on another compound produced by fireflies, firefly luciferin. In the presence of magnesium ions, ATP, and oxygen, this reaction produces light.[13][14] Oxygen is supplied via an abdominal trachea or breathing tube. Gene coding for these substances has been inserted into many different organisms.[15] Firefly luciferase is used in forensics, and the enzyme has medical uses – in particular, for detecting the presence of ATP or magnesium.[11] Fireflies produce a "cold light", with no infrared or ultraviolet frequencies. The light may be yellow, green, or pale red, with wavelengths from 510 to 670 nanometers. Some species such as the dimly glowing "blue ghost" of the Eastern US may seem to emit blueish-white light from a distance and in low light conditions, but their glow is bright green when observed up close.[16] Their perceived blue tint may be due to the Purkinje effect.[17] During a study on the genome of Aquatica leii, scientists discovered two key genes are responsible for the formation, activation, and positioning of this firefly's light organ: Alabd-B and AlUnc-4.[18]
Adults emit light primarily for mate selection. Early larval bioluminescence was adopted in the phylogeny of adult fireflies, and was repeatedly gained and lost before becoming fixed and retained as a mechanism of sexual communication in many species.[11][19] Adult lampyrids have a variety of ways to communicate with mates in courtships: steady glows, flashing, and the use of chemical signals unrelated to photic systems.[20] Chemical signals, or pheromones, are the ancestral form of sexual communication; this pre-dates the evolution of flash signaling in the lineage, and is retained today in diurnally-active species.[11][21] Some species, especially lightning bugs of the genera Photinus, Photuris, and Pyractomena, are distinguished by the unique courtship flash patterns emitted by flying males in search of females. In general, females of the genus Photinus do not fly, but do give a flash response to males of their own species. Signals, whether photic or chemical, allow fireflies to identify mates of their own species. Flash signaling characteristics include differences in duration, timing, color, number and rate of repetitions, height of flight, and direction of flight (e.g. climbing or diving) and vary interspecifically and geographically.[22]Template:Sfn When flash signals are not sufficiently distinguished between species in a population, sexual selection encourages divergence of signaling patterns.[22]
Synchronization of flashing occurs in several species; it is explained as phase synchronization and spontaneous order.[23] Tropical fireflies routinely synchronise their flashes among large groups, particularly in Southeast Asia. At night along river banks in the Malaysian jungles, fireflies synchronize their light emissions precisely. Hypotheses for the causes of this behavior involve diet, social interaction, and altitude. In the Philippines, thousands of fireflies can be seen all year-round in the town of Donsol. In the United States, one of the most famous sightings of fireflies blinking in unison occurs annually near Elkmont, Tennessee, in the Great Smoky Mountains during the first weeks of June.[24] Congaree National Park in South Carolina is another host to this phenomenon.[25]
Female "femme fatale" Photuris fireflies mimic the photic signaling patterns of the smaller Photinus, attracting males to what appears to be a suitable mate, then eating them.[11] This provides the females with a supply of the toxic defensive lucibufagin chemicals.Template:Sfn
Many fireflies do not produce light. Usually these species are diurnal, or day-flying, such as those in the genus Ellychnia. A few diurnal fireflies that inhabit primarily shadowy places, such as beneath tall plants or trees, are luminescent. One such genus is Lucidota.
Non-bioluminescent fireflies use pheromones to signal mates. Some basal groups lack bioluminescence and use chemical signaling instead. Phosphaenus hemipterus has photic organs, yet is a diurnal firefly and displays large antennae and small eyes. These traits suggest that pheromones are used for sexual selection, while photic organs are used for warning signals. In controlled experiments, males coming from downwind arrived at females first, indicating that males travel upwind along a pheromone plume. Males can find females without the use of visual cues, so sexual communication in P. hemipterus appears to be mediated entirely by pheromones.[26]
-
Lamprohiza female by her own light
-
A video of fireflies
-
Fireflies in the woods near Nuremberg, Germany, 30-second exposure
Evolution
Fossil history
The oldest known fossils of the Lampyridae family are Protoluciola and Flammarionella from the Late Cretaceous (Cenomanian ~ 99 million years ago) Burmese amber of Myanmar, which belong to the subfamily Luciolinae. The light producing organs are clearly present.[27][28] The ancestral glow colour for the last common ancestor of all living fireflies has been inferred to be green, based on genomic analysis.[29]
Taxonomy
The fireflies (including the lightning bugs) are a family, Lampyridae, of some 2,000 species within the Coleoptera. The family forms a single clade, a natural phylogenetic group.[30] The term glowworm is used for both adults and larvae of firefly species such as Lampyris noctiluca, the common European glowworm, in which only the nonflying adult females glow brightly; the flying males glow weakly and intermittently.[31][32][33] In the Americas, "glow worms" are the closely related Coleopteran family Phengodidae, while in New Zealand and Australia, a "glow worm" is a luminescent larva of the fungus gnat Arachnocampa, within the true flies, Diptera.[31]
Phylogeny
The phylogeny of the Lampyridae family, based on both phylogenetic and morphological evidence by Martin et al. 2019, is:[30]
Interaction with humans
Conservation
Firefly populations are thought to be declining worldwide. While monitoring data for many regions are scarce, a growing number of anecdotal reports, coupled with several published studies from Europe and Asia, suggest that fireflies are endangered.[34][35][36][37] Recent IUCN Red List assessments for North American fireflies have identified species with heightened extinction risk in the US, with 18 taxa categorized as threatened with extinction.[38][39]
Fireflies face threats including habitat loss and degradation, light pollution, pesticide use, poor water quality, invasive species, over-collection, and climate change.[40] Firefly tourism, a quickly growing sector of the travel and tourism industry, has also been identified as a potential threat to fireflies and their habitats when not managed appropriately.[41] Like many other organisms, fireflies are directly affected by land-use change (e.g., loss of habitat area and connectivity), which is identified as the main driver of biodiversity changes in terrestrial ecosystems.[42] Pesticides, including insecticides and herbicides, have also been indicated as a likely cause of firefly decline.[43][44] These chemicals can not only harm fireflies directly but also potentially reduce prey populations and degrade habitat. Light pollution is an especially concerning threat to fireflies. Since the majority of firefly species use bioluminescent courtship signals,[45] they are also sensitive to environmental levels of light and consequently to light pollution.[45][46] A growing number of studies investigating the effects of artificial light at night on fireflies has shown that light pollution can disrupt fireflies' courtship signals and even interfere with larval dispersal.[47][48][49][50] Researchers agree that protecting and enhancing firefly habitat is necessary to conserve their populations. Recommendations include reducing or limiting artificial light at night, restoring habitats where threatened species occur, and eliminating unnecessary pesticide use, among many others.[44][51][52]
In culture
Template:Multiple image Fireflies have featured in human culture around the world for centuries.[53] In Japan, the emergence of fireflies (Japanese: Template:Transliteration) signifies the anticipated changing of the seasons;[54] firefly viewing is a special aesthetic pleasure of midsummer, celebrated in parks that exist for that one purpose.[55] The Japanese sword Hotarumaru, made in the 14th century, is so named for a legend that its flaws were repaired by fireflies.[56][57]
In Italy, the firefly (Italian: Script error: No such module "Lang".) appears in Canto XXVI of Dante's Inferno, written in the 14th century:[58]
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Quante 'l villan ch'al poggio si riposa,
nel tempo che colui che 'l mondo schiara
la faccia sua a noi tien meno ascosa,
come la mosca cede a la zanzara,
vede lucciole giù per la vallea,
forse colà dov' e' vendemmia e ara:
di tante fiamme tutta risplendea
l'ottava bolgia, ...
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As many as the fireflies which the peasant sees in the [Tuscan] valley below, when he is resting on the hill—in the season [midsummer] when the sun hides least from us, and at the time of day [dusk] when the fly gives place to the mosquito—perhaps in the fields where he tills the ground and gathers in the grapes; with that many flames the eighth ditch [of Hell] was shining, ...
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References
Sources
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Further reading
- Faust, Lynn Frierson (2017). "Fireflies, Glow-worms, and Lightning Bugs"
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- Lewis, Sara (2016). Silent Sparks: The Wondrous World of Fireflies. Princeton University Press. Template:ISBN
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External links
Template:Sister project Template:Sister project
- An introduction to European fireflies and glow-worms
- Firefly.org – Firefly & Lightning Bug Facts, Pictures, Information About Firefly Insect Disappearance
- Firefly simulating robot, China
- NCBI taxonomy database
- Museum of Science, Boston – Understanding Fireflies
- Video of a firefly larva in Austria
- FireflyExperience.org – Luminous Photography and Videos of Fireflies & Lightning Bugs
Template:Coleoptera Template:Taxonbar Template:Authority control
- ↑ Script error: No such module "citation/CS1".
- ↑ BugGuide: common names
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- ↑ Synchronous Fireflies – Great Smoky Mountains National Park. Nps.gov (3 June 2013). Retrieved on 22 June 2013.
- ↑ Cross, Robert (23 May 2004) "Tree huggin'". Chicago Tribune.
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- ↑ Potter, Daniel A. See "How You Can Help", FireFly.org, citing (1) "Understanding Halofenozide (Mach 2) and Imidacloprid (Merit) Soil Insecticides," International SportsTurf Institute, Incorporated, Turfax, Vol. 6, No. 1 (Jan-Feb 1998) and (2) "Relative Toxicities of Chemicals to the Earthworm Eisenia foetida," by Brian L. Roberts and H. Wyman Dorough. Article first published online: 20 October 2009. Environmental Toxicology and Chemistry, Vol. 3, No. 1 (Jan. 1984), pp. 67–78.
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