Firefly: Difference between revisions

Firefly
	Unidentified species from India, dorsal (left) and ventral aspect
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Infraclass:	Neoptera
Superorder:	Endopterygota
Order:	Coleoptera
Suborder:	Polyphaga
Infraorder:	Elateriformia
Superfamily:	Elateroidea
Family:	Lampyridae; Latreille, 1817
Subfamilies
	Cyphonocerinae; Lampyrinae; Luciolinae; Ototetrinae (disputed); Photurinae; and see below Genera incertae sedis:; Oculogryphus; Pterotus LeConte, 1859

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Lampyridae is a family in the beetle order Coleoptera. They are commonly called fireflies, lightning bugs due to their conspicuous nocturnal or more accurately crepuscular use of bioluminescence to attract mates or prey. Fireflies are capable of producing a "cold light" containing no ultraviolet or infrared rays, with a wavelength from 510 to 670 nanometers, pale reddish, yellowish or green in colour, with a lighting efficiency of up to 96%^{[citation needed]}.

There are more than 2,000 species of firefly found in temperate and tropical environments around the world. Many species can be found in marshes or in wet, wooded areas where their larvae have abundant sources of food. These can also emit light and are often called "glowworms", particularly in Eurasia. In the Americas, "glow worm" also refers to the related Phengodidae.

Biology

Fireflies tend to be brown and soft-bodied, often with the elytra more leathery than in other beetles. Though the females of some species are similar in appearance to males, larviform females are found in many other firefly species. These females can often be distinguished from the larvae only because they have compound eyes. The most commonly known fireflies are nocturnal, though there are numerous species that are diurnal. Most diurnal species are non-luminescent, though some species that remain in shadowy areas can produce light.

A few days after mating, a female lays her fertilized eggs on or just below the surface of the ground. The eggs hatch 3-4 weeks later and the larva feed until the end of the summer. The larvae are commonly called glowworms, not to be confused with the distinct beetle family Phengodidae or fly genus Arachnocampa. Lampyrid larvae have simple eyes. The term glowworm is also used for both adults and larvae of species such as Lampyris noctiluca, the common European glowworm, in which only the non-flying adult females glow brightly and the flying males glow only very weakly and intermittently. Fireflies overwinter (some species for several years) during the larval stage. Some do this by burrowing underground, while others find places on or under the bark of trees. They emerge in the spring. After several weeks of feeding, they pupate for 1 to 2.5 weeks and emerge as adults. 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 which deliver digestive fluids directly to their prey. The diet of adults is variable. It has been reported that some are predatory, while others find food on plant pollen and sometimes nectar.

Light production

Fireflies in the woods near Nuremberg, Germany. Exposure time 30 seconds.

Light production in fireflies is due to a type of chemical reaction called bioluminescence, which, in fireflies, occurs in specialized light-emitting organs usually on the lower abdomen. The enzyme luciferase acts on luciferin in this organ to stimulate light emission. Genes coding for these substances have been inserted into many different organisms (see Luciferase - Applications). Luciferase is also used in forensics, and the enzyme has medical uses.

For adult beetles, it is primarily used to locate other individuals of the same species for reproduction. Many species, especially the famous 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. Photinus females as usual in this family generally do not fly, but give a flash response to males of their own species.

Bioluminescence is a very efficient process. Some 90% of the energy a firefly uses to create light is actually converted into visible light^{[citation needed]}. By comparison, an incandescent electric bulb can convert only 10 percent of total energy used into visible light, and the remainder is emitted as heat.

Tropical fireflies, particularly in Southeast Asia (Thailand and Malaysia), routinely synchronize their flashes among large groups, a startling example of spontaneous biological order. This phenomenon occurs through the night along river banks in the Malaysian jungles every day of the year. Current hypotheses about the causes of this behavior involve diet, social interaction, and altitude. In the United States, one of the most famous sightings of fireflies blinking in unison occurred near Elkmont, Tennessee in the Great Smoky Mountains during the second week of June 2005[1]. Congaree National Park in South Carolina is another host to the phenomenon [2].

Female Photuris fireflies are known for mimicking the mating flashes of other "lightning bugs" for the sole purpose of predation. Target males are attracted to what appears to be a suitable mate, and are then eaten. For this reason the Photuris species are sometimes referred to as "femme fatale fireflies".

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 primarily inhabit shadowy places, such as beneath tall plants or trees, are luminescent. One such genus is Lucidota.

All fireflies glow as larvae. Bioluminescence serves a different function in lampyrid larvae than it does in adults. It appears to be a warning signal to predators, since many firefly larvae contain chemicals that are distasteful or toxic.

Systematics

Firefly systematics, as with many insects, are in a constant state of flux, as new species continue to be discovered. The five subfamilies listed above are the most commonly accepted ones, though others such as the Amydetinae and Psilocladinae have been proposed. This was mainly done in an attempt to revise the Lampyrinae, which by and by had become something of a "wastebin taxon" to hold incertae sedis species and genera of fireflies. Other changes have been proposed, such as merging the Ototetrinae into the Luciolinae, but the arrangement used here appears to be the most frequently-seen and stable layout for the time being. Though most groups by and large appear to be monophyletic, some (e.g. the tribe Photinini) are perhaps better split up.

There seem to be two groups of subfamilies: one containing many American and some Eurasian species in the Lampyrinae and Photurinae, and one predominantly Asian one made up from the other subfamilies. While the subfamilies as understood here are monophyletic by and large, there are still a few genera that need to be moved about for the subfamilies to accurately represent the evolutionary relationships among the fireflies.

The Rhagophthalmidae are a glow worm-like lineage of Elateroidea. They have in the recent past usually been consiered a distinct family, but it is still disputed whether this is correct. Indeed, they might be the only close relative of the puzzling firefly genus Pterotus which sometimes is placed in a monotypic subfamily.

The genus Phausis, usually placed in the tribe Photinini of the Lampyrinae, might represent another rather distinct lineage instead.

Fireflies and humans

Fireflies were a part of ancient Mayan mythology, often being associated with the stars. Further, they were associated with cigar smoking and may have had at least one representative in the pantheon of Mayan gods.^[1]

In East Asia, the ancient Chinese sometimes captured fireflies in transparent or semi-transparent containers and used them as (short-term) lanterns^{[citation needed]}. In Philippine folklore, a tree surrounded by fireflies is believed to be haunted by a tikbalang, the local term for a mischievous satyr.

The Pennsylvania Firefly (Photuris pennsylvanica) is the state insect of Pennsylvania, and the Common Eastern Firefly (P. pyralis) one of the state insects of Tennessee. At one point, Indiana seriously considered making the State's insect a firefly, but the legislature never put the measure to a vote.

The spectacular synchronized flashing by Pteroptyx and other Luciolinae fireflies has potential economic significance. Notably on the Selangor River at Kampong Kuantan (close to Kuala Selangor, Malaysia), it has become a major attraction for tourists, creating considerable revenue for the local economy.

Anecdotal evidence suggests human activities may be impacting firefly populations negatively, with urban sprawl and light pollution cited as potential culprits.^[2]

References

^ Lopes, Luís. 2004. "Some notes on fireflies." Mesoweb
^ Casey, Michael. "Experts fear firefly populations are blinking out." USA Today 2 Sep. 2008. Web.

http://members.jcom.home.ne.jp/hotaru-net/. A site about Japanese aquatic firefly habits, life-history, biology, resources, and activities.
http://members.jcom.home.ne.jp/hotaru-net/photo/firefly.html
http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi
http://www.iisc.ernet.in/academy/resonance/Sept2002/pdf/Sept2002p49-55.pdf
Branham, M. A., and J. W. Wenzel. 2003. The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae). Cladistics 19: 1-22.
Stous, Hollend. 1997. A review of predation in Photuris, and its effects on the evolution of flash signaling in other New World fireflies. http://www.colostate.edu/Depts/Entomology/courses/en507/papers_1997/stous.html
http://www.fireflies.tk . A site about the Firefly meeting 2007 in Portugal and information on fireflies in general.
http://pirilampos-lightalive.blogspot.com/. A site about bioluminescence and Firefly project in Portugal. Funny

[lopes-1] Lopes, Luís. 2004. "Some notes on fireflies." Mesoweb

[decline-2] Casey, Michael. "Experts fear firefly populations are blinking out." USA Today 2 Sep. 2008. Web.

[1]

[2]

@@ Line 2: / Line 2: @@
 {{Taxobox
 | name = Firefly
-| image = Photuris_lucicrescens.jpg
+| image = Firefly photuris.jpg
 | image_width = 240px
-| image_caption = Adult ''[[Photuris lucicrescens]]'' firefly
+| image_caption = Unidentified species from [[India]], dorsal (left) and ventral aspect
 | regnum = [[Animal]]ia
 | phylum = [[Arthropod]]a
@@ Line 21: / Line 21: @@
 [[Lampyrinae]]<br />
 [[Luciolinae]]<br />
-[[Ototetrinae]]<br />
+[[Ototetrinae]] <small>(disputed)</small><br />
 [[Photurinae]]<br />
 and see [[#Systematics|below]]
 ----
 [[Genera]] ''[[incertae sedis]]'':<br />
-''[[Oculogryphus]]''<!-- AmMusNovit3600 --><br />
+''[[Oculogryphus]]''<br /><!-- AmMusNovit3600 -->
-''[[Pterotus]]''
+''[[Pterotus]]'' <small>LeConte, 1859</small><!-- MolPhylEvol45:22. -->
 }}
-'''Lampyridae''' is a [[family (biology)|family]] in the [[beetle]] [[order (biology)|order]] Coleoptera, members of which are commonly called '''fireflies''', '''lightning bugs''' or (in their [[larva]]l state) [[glow worm]]s due to their conspicuous [[Nocturnal animal|nocturnal]] (or, more accurately, [[crepuscular]]) use of [[bioluminescence]] to attract mates or prey. Fireflies are capable of producing a "cold light" containing no [[ultraviolet]] or [[infrared]] rays, with a wavelength from 510 to 670 [[nanometer]]s, pale reddish, yellowish or green in colour, with a [[lighting efficiency]] of up to 96%{{Fact|date=May 2008}}.
+'''Lampyridae''' is a [[family (biology)|family]] in the [[beetle]] [[order (biology)|order]] Coleoptera. They are commonly called '''fireflies''', '''[[lightning bug]]s''' due to their conspicuous [[Nocturnal animal|nocturnal]] or more accurately [[crepuscular]] use of [[bioluminescence]] to attract mates or prey. Fireflies are capable of producing a "cold light" containing no [[ultraviolet]] or [[infrared]] rays, with a wavelength from 510 to 670 [[nanometer]]s, pale reddish, yellowish or green in colour, with a [[lighting efficiency]] of up to 96%{{Fact|date=May 2008}}.
-There are more than 2000 [[species]] of firefly found in [[temperate]] and [[tropical]] environments around the world. Many species can be found in marshes or in wet, wooded areas where their [[larva]]e have abundant sources of food.
+There are more than 2,000 [[species]] of firefly found in [[temperate]] and [[tropical]] environments around the world. Many species can be found in marshes or in wet, wooded areas where their [[larva]]e have abundant sources of food. These can also emit light and are often called '''"[[glowworm]]s"''', particularly in Eurasia. In the [[Americas]], "glow worm" also refers to the related [[Phengodidae]].
 ==Biology==
-[[Image:Lampyridae.jpg|left|thumb|[[Scanning electron microscope|Scanning electron micrograph]] of a firefly]]
+[[Image:Luciola4 crop.png|left|thumb|A [[larviform female]] showing light-emitting organs on [[abdomen]]]]
 Fireflies tend to be brown and soft-bodied, often with the [[elytra]] more leathery than in other beetles. Though the females of some species are similar in appearance to males, [[larviform female]]s are found in many other firefly species. These females can often be distinguished from the larvae only because they have [[compound eye]]s. The most commonly known fireflies are nocturnal, though there are numerous species that are [[diurnal animal|diurnal]]. Most diurnal species are non-luminescent, though some species that remain in shadowy areas can produce light.
@@ Line 41: / Line 41: @@
 ===Light production===
-[[Image:Firefly composite.jpg|thumb|right|Firefly (species unknown) captured in Eastern Canada. The top picture is taken with a flash, the bottom only with the self-emitted light.]]
+[[Image:Firefly composite.jpg|thumb|left|Firefly (species unknown) captured in Eastern Canada. The top picture is taken with a flash, the bottom only with the self-emitted light.]]
+[[Image:GluehwuermchenImWald.jpg|thumb|right|Fireflies in the woods near [[Nuremberg, Germany]]. Exposure time 30 seconds.]]
-[[Image:GluehwuermchenImWald.jpg|thumb|right|Fireflies in the woods near [[Nuremberg, Germany]]]]
 Light production in fireflies is due to a type of chemical reaction called [[bioluminescence]], which, in fireflies, occurs in specialized light-emitting [[organ (biology)|organs]] usually on the lower abdomen.  The enzyme [[luciferase]] acts on [[luciferin]] in this organ to stimulate light emission. [[Gene]]s coding for these substances have been inserted into many different organisms (see [[Luciferase#Applications|Luciferase - Applications]]). Luciferase is also used in [[forensics]], and the enzyme has medical uses.
-For adult beetles, it is primarily used to locate other individuals of the same species for reproduction. Many species, especially in the genus ''[[Photinus (genus)|Photinus]]'', are distinguished by the unique courtship flash patterns emitted by flying males in search of females. ''Photinus'' females generally do not fly, but give a flash response to males of their own species.
+For adult beetles, it is primarily used to locate other individuals of the same species for reproduction. Many species, especially the famous '''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. ''Photinus'' females as usual in this family generally do not fly, but give a flash response to males of their own species.
 Bioluminescence is a very efficient process. Some 90% of the energy a firefly uses to create light is actually converted into visible light{{Fact|date=July 2008}}<!-- same citation also needed in article Luciferase -->. By comparison, an incandescent electric bulb can convert only 10 percent of total energy used into visible light, and the remainder is emitted as heat.
+[[Image:Leuchtkäfer - Firefly.JPG|thumb|right|Firefly larva]]
-Tropical fireflies, particularly in Southeast Asia ([[Thailand]] and [[Malaysia]]), routinely synchronize their flashes among large groups, a startling example of spontaneous biological order. This phenomenon occurs through the night along river banks in the Malaysian jungles every day of the year. Current hypotheses about the causes of this behavior involve diet, social interaction, and altitude. In the [[United States]], one of the most famous sightings of fireflies blinking in [[unison]] occurred near [[Elkmont, Tennessee]] in the [[Great Smoky Mountains]] during the second week of June 2005[[Image:Leuchtkäfer - Firefly.JPG|thumb|right|Firefly larva]][http://www.nps.gov/grsm/pphtml/newsdetail18318.html]. [[Congaree National Park]] in [[South Carolina]] is another host to the phenomenon [http://www.seacoastonline.com/2004news/05232004/travel/17745.htm].
+Tropical fireflies, particularly in Southeast Asia ([[Thailand]] and [[Malaysia]]), routinely synchronize their flashes among large groups, a startling example of spontaneous biological order. This phenomenon occurs through the night along river banks in the Malaysian jungles every day of the year. Current hypotheses about the causes of this behavior involve diet, social interaction, and altitude. In the [[United States]], one of the most famous sightings of fireflies blinking in [[unison]] occurred near [[Elkmont, Tennessee]] in the [[Great Smoky Mountains]] during the second week of June 2005[http://www.nps.gov/grsm/pphtml/newsdetail18318.html]. [[Congaree National Park]] in [[South Carolina]] is another host to the phenomenon [http://www.seacoastonline.com/2004news/05232004/travel/17745.htm].
-Female ''[[Photuris (genus)|Photuris]]'' fireflies are known for mimicking the mating flashes of other fireflies for the sole purpose of [[predator|predation]]. Target males are attracted to what appears to be a suitable mate, and are then eaten. For this reason the ''Photuris'' female is sometimes referred to as "[[femme fatale]]".
+Female ''[[Photuris (genus)|Photuris]]'' fireflies are known for mimicking the mating flashes of other "lightning bugs" for the sole purpose of [[predator|predation]]. Target males are attracted to what appears to be a suitable mate, and are then eaten. For this reason the ''Photuris'' species are sometimes referred to as "[[femme fatale]] fireflies".
 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 primarily inhabit shadowy places, such as beneath tall plants or trees, are luminescent. One such genus is ''[[Lucidota]]''.
@@ Line 60: / Line 59: @@
 ===Systematics===
+[[Image:Cyphonocerus ruficollis 2552543412 crop.png|thumb|left|''[[Cyphonocerus ruficollis]]'', a weakly-glowing member of the [[Cyphonocerinae]]]]
-Firefly systematics, as with many insects, are in a constant state of flux, as new species continue to be discovered. The five [[subfamilies]] listed above are the most commonly accepted ones, though others such as the [[Amydetinae]] and [[Psilocladinae]] have been proposed. This was mainly done in an attempt to revise the [[Lampyrinae]], which by and by had become something of a "[[wastebin taxon]]" to hold ''[[incertae sedis]]'' species and genera of fireflies. Other changes are occasionally proposed, such as merging the [[Ototetrinae]] into the [[Luciolinae]], but the arrangement used here appears to be the most frequently-seen and stable layout, at least for the time being.
+Firefly systematics, as with many insects, are in a constant state of flux, as new species continue to be discovered. The five [[subfamilies]] listed above are the most commonly accepted ones, though others such as the [[Amydetinae]] and [[Psilocladinae]] have been proposed. This was mainly done in an attempt to revise the [[Lampyrinae]], which by and by had become something of a "[[wastebin taxon]]" to hold ''[[incertae sedis]]'' species and genera of fireflies. Other changes have been proposed, such as merging the [[Ototetrinae]] into the [[Luciolinae]], but the arrangement used here appears to be the most frequently-seen and stable layout for the time being. Though most groups by and large appear to be [[monophyletic]], some (e.g. the [[tribe (biology)|tribe]] [[Photinini]]) are perhaps better split up.
+There seem to be two groups of subfamilies: one containing many [[America]]n and some [[Eurasia]]n species in the Lampyrinae and Photurinae, and one predominantly [[Asia]]n one made up from the other subfamilies. While the subfamilies as understood here are [[monophyletic]] by and large, there are still a few genera that need to be moved about for the subfamilies to accurately represent the [[evolution]]ary relationships among the fireflies.
+The [[Rhagophthalmidae]] are a [[Phengodidae|glow worm]]-like lineage of [[Elateroidea]]. They have in the recent past usually been consiered a distinct family, but it is still disputed whether this is correct. Indeed, they might be the only close relative of the puzzling firefly genus ''[[Pterotus]]'' which sometimes is placed in a [[monotypic]] subfamily.
+The genus ''[[Phausis]]'', usually placed in the [[tribe (biology)|tribe]] [[Photinini]] of the Lampyrinae, might represent another rather distinct lineage instead.
 ==Fireflies and humans==
@@ Line 68: / Line 74: @@
 In [[East Asia]], the ancient [[China|Chinese]] sometimes captured fireflies in transparent or semi-transparent containers and used them as (short-term) [[lantern]]s{{Fact|date=December 2007}}.  In [[Philippine folklore]], a tree surrounded by fireflies is believed to be haunted by a [[tikbalang]], the local term for a mischievous satyr.
-The [[Pennsylvania Firefly]] (''[[Photuris pennsylvanica]]'') is the [[state insect]] of [[Pennsylvania]], and the [[Common Eastern Firefly]] (''[[Photuris pyralis]]'') one of the state insects of [[Tennessee]]. At one point, [[Indiana]] seriously considered making the State's insect a firefly, but the legislature never put the measure to a vote.
+The [[Pennsylvania Firefly]] (''Photuris pennsylvanica'') is the [[state insect]] of [[Pennsylvania]], and the [[Common Eastern Firefly]] (''P. pyralis'') one of the state insects of [[Tennessee]]. At one point, [[Indiana]] seriously considered making the State's insect a firefly, but the legislature never put the measure to a vote.
 The spectacular synchronized flashing by ''[[Pteroptyx]]'' and other [[Luciolinae]] fireflies has potential economic significance. Notably on the [[Selangor River]] at [[Kampong Kuantan]] (close to  [[Kuala Selangor]], [[Malaysia]]), it has become a major attraction for tourists, creating considerable revenue for the local economy.
@@ Line 74: / Line 80: @@
 Anecdotal evidence suggests human activities may be impacting firefly populations negatively, with urban sprawl and light pollution cited as potential culprits.<ref name="decline">Casey, Michael. "Experts fear firefly populations are blinking out." [http://www.usatoday.com/tech/science/2008-09-02-firefly-population-endangered_N.htm USA Today 2 Sep. 2008]. Web.</ref>
-==References==
+==References==<!-- MolPhylEvol45:22. -->
 {{Commonscat|Lampyridae}}
 {{reflist}}
-<!--* {{ITIS|ID=113835|taxon=Lampyridae|date=30 April|year=2006}} [subfamily list is erroneous, see AmMusNovit3600 -->
+<!--* {{ITIS|ID=113835|taxon=Lampyridae|date=30 April|year=2006}} [subfamily list is erroneous, see AmMusNovit3600] -->
 * http://members.jcom.home.ne.jp/hotaru-net/. A site about Japanese aquatic firefly habits, life-history, biology, resources, and activities.
 * http://members.jcom.home.ne.jp/hotaru-net/photo/firefly.html
@@ Line 88: / Line 94: @@
 [[Category:Bioluminescent organisms]]
-[[Category:Lampyridae|*]]
+[[Category:Lampyridae| ]]
 [[Category:Night]]
 [[Category:Insect families]]