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{{Use dmy dates|date=August 2013}}
{{short description|Family of butterflies}}
{{short description|Family of butterflies}}
{{Use dmy dates|date=November 2021}}
{{Automatic taxobox
{{Automatic taxobox
| image = Maculinea arion Large Blue Upperside SFrance 2009-07-18.jpg
| image = Maculinea arion Large Blue Upperside SFrance 2009-07-18.jpg
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and see [[#Subfamilies|text]] or [[List of lycaenid genera]]
and see [[#Subfamilies|text]] or [[List of lycaenid genera]]
}}
}}
[[File:Brown hairstreak (Thecla betulae) Bulgaria.jpg|thumb|Brown hairstreak (''[[Thecla betulae]]'')]]

[[File:Loxura atymnus-Kadavoor-2017-04-23-001.jpg|thumb|''[[Loxura atymnus]]'']]
'''Lycaenidae''' is the second-largest [[family (biology)|family]] of [[butterflies]] (behind [[Nymphalidae]], brush-footed butterflies), with over 6,000 species worldwide,<ref>Pierce, Naomi E.; Braby, Michael F.; Heath, Alan; Lohman, David J.; Mathew, John; Rand, Douglas B. & Travassos, Mark A. (January 2002).[http://www.annualreviews.org/doi/pdf/10.1146/annurev.ento.47.091201.145257 "The Ecology and Evolution of Ant Association in the Lycaenidae (Lepidoptera)". ''Annual Review of Entomology''. '''47''' (1): 259-267. {{doi|10.1146/annurev.ento.47.091201.145257}}.</ref> whose members are also called '''gossamer-winged butterflies'''. They constitute about 30% of the known butterfly species.
'''Lycaenidae''' is the second-largest [[family (biology)|family]] of [[butterflies]] (behind [[Nymphalidae]], brush-footed butterflies), with over 6,000 species worldwide,<ref>{{cite journal |last1=Pierce |first1=Naomi E. |last2=Braby |first2=Michael F. |last3=Heath |first3=Alan |last4=Lohman |first4=David J. |last5=Mathew |first5=John |last6=Rand |first6=Douglas B. |last7=Travassos |first7=Mark A. |title=The Ecology and Evolution of Ant Association in the Lycaenidae (Lepidoptera) |journal=Annual Review of Entomology |date=January 2002 |volume=47 |issue=1 |pages=733–771 |doi=10.1146/annurev.ento.47.091201.145257 |pmid=11729090 }}</ref> whose members are also called '''gossamer-winged butterflies'''. They constitute about 30% of the known butterfly species.


The family comprises seven subfamilies, including the blues ([[Polyommatinae]]), the coppers ([[Lycaeninae]]), the hairstreaks ([[Theclinae]]), and the harvesters ([[Miletinae]]).
The family comprises seven subfamilies, including the blues ([[Polyommatinae]]), the coppers ([[Lycaeninae]]), the hairstreaks ([[Theclinae]]), and the harvesters ([[Miletinae]]).


==Description, food, and life cycle==
==Description, food, and life cycle==
Adults are small, under 5&nbsp;cm usually, and brightly coloured, sometimes with a metallic gloss. Lycaenidae wings are generally blue or green. More than half of these butterflies depend on ants in some way.<ref>{{cite journal |last1=Ueda |first1=Shouhei |last2=Komatsu |first2=Takashi |last3=Itino |first3=Takao |last4=Arai |first4=Ryusuke |last5=Sakamoto |first5=Hironori |title=Host-ant specificity of endangered large blue butterflies (Phengaris spp., Lepidoptera: Lycaenidae) in Japan |journal=Scientific Reports |date=3 November 2016 |volume=6 |issue=1 |pages=36364 |doi=10.1038/srep36364 |pmid=27808223 |pmc=5093462 |bibcode=2016NatSR...636364U }}</ref>
Adults are small, under 5&nbsp;cm usually, and brightly coloured, sometimes with a metallic gloss.


Larvae are often flattened rather than cylindrical, with glands that may produce secretions that attract and subdue [[ants]]. Their cuticles tend to be thickened. Some larvae are capable of producing vibrations and low sounds that are transmitted through the substrates they inhabit. They use these sounds to communicate with ants.<ref name="pierce">Pierce, N. E.; Braby, M. F.; Heath, A.; Lohman, D. J.; Mathew, J.; Rand, D. B. & Travassos, M. A. (2002). [http://www.oeb.harvard.edu/faculty/pierce/publications/pdfs/2002_Pierce_et_al.pdf "The ecology and evolution of ant association in the Lycaenidae (Lepidoptera)"]. ''Annual Review of Entomology''. '''47''': 733-771. </ref><ref>DeVries, Philip J. (1992). "Singing Caterpillars, Ants and Symbiosis". ''Scientific American'', '''267''': 76</ref>
Larvae are often flattened rather than cylindrical, with glands that may produce secretions that attract and subdue [[ants]]. Their cuticles tend to be thickened. Some larvae are capable of producing vibrations and low sounds that are transmitted through the substrates they inhabit. They use these sounds to communicate with ants.<ref name="pierce">{{cite journal |last1=Pierce |first1=Naomi E. |last2=Braby |first2=Michael F. |last3=Heath |first3=Alan |last4=Lohman |first4=David J. |last5=Mathew |first5=John |last6=Rand |first6=Douglas B. |last7=Travassos |first7=Mark A. |title=The Ecology and Evolution of Ant Association in the Lycaenidae (Lepidoptera) |journal=Annual Review of Entomology |date=January 2002 |volume=47 |issue=1 |pages=733–771 |id={{ProQuest|222345887}} |doi=10.1146/annurev.ento.47.091201.145257 |pmid=11729090 }}</ref><ref>{{cite journal |last1=DeVries |first1=Philip J. |title=Singing Caterpillars, Ants and Symbiosis |journal=Scientific American |date=1992 |volume=267 |issue=4 |pages=76–83 |doi=10.1038/scientificamerican1092-76 |jstor=24939256 |bibcode=1992SciAm.267d..76D }}</ref>


Adult individuals often have hairy antenna-like tails complete with black and white annulated (ringed) appearance. Many species also have a spot at the base of the tail and some turn around upon landing to confuse potential predators from recognizing the true head orientation. This causes predators to approach from the true head end resulting in early visual detection.<ref>Robbins, Robert K. (1981). "The 'False Head' Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies". ''American Naturalist''. '''118''' (5): 770-775.</ref>
Adult individuals often have hairy antenna-like tails complete with black and white annulated (ringed) appearance. Many species also have a spot at the base of the tail and some turn around upon landing to confuse potential predators from recognizing the true head orientation. This causes predators to approach from the true head end resulting in early visual detection or to attack the false head ending up with a beak of dusty scales.<ref>{{cite journal |last1=Robbins |first1=Robert K. |title=The 'False Head' Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies |journal=The American Naturalist |date=1981 |volume=118 |issue=5 |pages=770–775 |doi=10.1086/283868 |jstor=2460614 |s2cid=34146954 }}</ref>


Lycaenid caterpillars are diverse in their food habits and apart from [[phytophagy]], some are [[entomophagous]], feeding on [[aphids]], [[scale insects]], and ant larvae. Some lycaenids even exploit their association with ants by inducing ants to feed them by regurgitation, a process called [[trophallaxis]]. Not all lycaenid butterflies need ants, but about 75% of species associate with ants,<ref name="pierce"/> a relationship called [[myrmecophily]]. These associations can be mutualistic, parasitic, or predatory depending on the species.
[[File:Lycaenid ant sec.jpg|thumb|left|Ant tending a lycaenid larva]]
Lycaenids are diverse in their food habits and apart from [[phytophagy]], some of them are [[entomophagous]], feeding on [[aphids]], [[scale insects]], and ant larvae. Some lycaenids even exploit their association with ants by inducing ants to feed them by regurgitation, a process called [[trophallaxis]]. Not all lycaenid butterflies need ants, but about 75% of species associate with ants,<ref name="pierce"/> a relationship called [[myrmecophily]]. These associations can be mutualistic, parasitic, or predatory depending on the species.


In some species, larvae are attended and protected by ants while feeding on the host plant, and the ants receive sugar-rich [[Honeydew (secretion)|honeydew]] from them, throughout the larval life, and in some species during the pupal stage. In other species, only the first few [[instar]]s are spent on the plant, and the remainder of the larval lifespan is spent as a predator within the ant nest. It becomes a parasite, feeding on ant regurgitations, or a predator on the ant larvae.<ref name="pierce"/> The caterpillars pupate inside the ants' nest and the ants continue to look after the pupae. Just before the adults emerge, the wings of the butterfly inside the pupal case detach from it, and the pupa becomes silvery. The adult butterfly emerges from the pupa after three to four weeks, still inside the ant nest. The butterfly must crawl out of the ant nest before it can expand its wings.
In some species, larvae are attended and protected by ants while feeding on the host plant, and the ants receive sugar-rich [[Honeydew (secretion)|honeydew]] from them, throughout the larval life, and in some species during the pupal stage. In other species, only the first few [[instar]]s are spent on the plant, and the remainder of the larval lifespan is spent as a predator within the ant nest. It becomes a parasite, feeding on ant regurgitations, or a predator on the ant larvae.<ref name="pierce"/> The caterpillars pupate inside the ants' nest and the ants continue to look after the pupae. Just before the adults emerge, the wings of the butterfly inside the pupal case detach from it, and the pupa becomes silvery. The adult butterfly emerges from the pupa after three to four weeks, still inside the ant nest. The butterfly must crawl out of the ant nest before it can expand its wings.
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==Subfamilies==
==Subfamilies==
[[File:Avene fg04.JPG|thumb|Mating ''[[Cyaniris semiargus]]'']]
[[File:Avene fg04.JPG|thumb|Mating ''[[Cyaniris semiargus]]'']]

[[File:Eumaeus atalaPCCP20040417-4033A.jpg|thumb|Caterpillar of the [[Eumaeus atala|Atala]] (''Eumaeus atala'')]]
[[File:Leaf Blue (Amblypodia anita) W IMG 2700.jpg|thumb|Leaf blue (''[[Amblypodia anita]]'') from [[Theclinae]]]]
Many taxonomists only include the [[Lycaeninae]], [[Theclinae]], [[Polyommatinae]], [[Poritiinae]], [[Miletinae]], and [[Curetinae]] under the Lycaenidae.<ref name="tolweb.org">Brower, Andrew V. Z. (2008). [http://tolweb.org/Lycaenidae/12175 "Lycaenidae [Leach&#93; 1815"]. Version 25 April 2008 (under construction). ''The Tree of Life Web Project''.</ref><ref>Ackery, P. R.; de Jong, R. & Vane-Wright, R. I. (1999). "The butterflies: Hedyloidea, Hesperioidea, and Papilionoidea". Pages 264-300 in: ''Lepidoptera: Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. Handbook of Zoology'' Vol. IV, Part 35. N. P. Kristensen, ed. De Gruyter, Berlin and New York.</ref>
Many taxonomists only include the [[Lycaeninae]], [[Theclinae]], [[Polyommatinae]], [[Poritiinae]], [[Miletinae]], and [[Curetinae]] under the Lycaenidae.<ref name="tolweb.org">Brower, Andrew V. Z. (2008). [http://tolweb.org/Lycaenidae/12175 "Lycaenidae [Leach&#93; 1815"]. Version 25 April 2008 (under construction). ''The Tree of Life Web Project''.</ref><ref>Ackery, P. R.; de Jong, R. & Vane-Wright, R. I. (1999). "The butterflies: Hedyloidea, Hesperioidea, and Papilionoidea". Pages 264-300 in: ''Lepidoptera: Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. Handbook of Zoology'' Vol. IV, Part 35. N. P. Kristensen, ed. De Gruyter, Berlin and New York.</ref>
The [[Aphnaeinae]], which used to be a tribe (Aphnaeini) within the Theclinae, were recently given subfamily rank too.<ref name="Boyle et al., 2015">Boyle, J. H.; Kaliszewska, Z. A.; Espeland, M.; Suderman, T. R.; Fleming, J.; Heath, A. & Pierce, N. E. (2015). "Phylogeny of the Aphnaeinae: Myrmecophilous African butterflies with carnivorous and herbivorous life histories". ''Systematic Entomology''. '''40''' (1): 169–182.</ref>
The [[Aphnaeinae]], which used to be a tribe (Aphnaeini) within the Theclinae, were recently given subfamily rank too.<ref name="Boyle et al., 2015">{{cite journal |last1=Boyle |first1=John H. |last2=Kaliszewska |first2=Zofia A. |last3=Espeland |first3=Marianne |last4=Suderman |first4=Tamara R. |last5=Fleming |first5=Jake |last6=Heath |first6=Alan |last7=Pierce |first7=Naomi E. |title=Phylogeny of the A phnaeinae: myrmecophilous A frican butterflies with carnivorous and herbivorous life histories |journal=Systematic Entomology |date=January 2015 |volume=40 |issue=1 |pages=169–182 |doi=10.1111/syen.12098 |bibcode=2015SysEn..40..169B |s2cid=83052979 }}</ref>


* [[Curetinae]] – sunbeams (Oriental or Palaearctic). Selected species:
* [[Curetinae]] – sunbeams (Oriental or Palaearctic). Selected species:
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* [[Aphnaeinae]] (Afrotropical and Oriental)
* [[Aphnaeinae]] (Afrotropical and Oriental)
* [[Theclinae]] – hairstreaks (usually tailed) and elfins (not tailed) (global). Selected species:
* [[Theclinae]] – hairstreaks (usually tailed) and elfins (not tailed) (global). Selected species:
** ''[[Arhopala]]'' - oakblues
** ''[[Arhopala]]'' oakblues
** ''[[Atlides halesus]]'' – great purple hairstreak
** ''[[Atlides halesus]]'' – great purple hairstreak
** ''[[Eumaeus atala]]'' – Atala
** ''[[Eumaeus atala]]'' – Atala
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** ''[[Lycaena dispar]]'' – large copper
** ''[[Lycaena dispar]]'' – large copper
** ''[[Lycaena phlaeas]]'' – small copper
** ''[[Lycaena phlaeas]]'' – small copper
** ''[[Lycaena heteronea]]'' - blue copper
** ''[[Lycaena heteronea]]'' blue copper
* [[Polyommatinae]] – blues (global). Selected species:
* [[Polyommatinae]] – blues (global). Selected species:
** ''[[Celastrina ladon]]'' – spring azure
** ''[[Celastrina ladon]]'' – spring azure
** ''[[Chilades]]'' - jewel blues
** ''[[Chilades]]'' jewel blues
** ''[[Cupido comyntas]]'' – eastern tailed-blue
** ''[[Cupido comyntas]]'' – eastern tailed-blue
** ''[[Cupido minimus]]'' – small blue
** ''[[Cupido minimus]]'' – small blue
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== References ==
== References ==
{{Reflist|2}}
{{reflist}}


== Further reading ==
== Further reading ==
Line 105: Line 104:
{{Wikispecies}}
{{Wikispecies}}
* [http://tolweb.org/tree?group=Lycaenidae ''Tree of Life Web Project'']
* [http://tolweb.org/tree?group=Lycaenidae ''Tree of Life Web Project'']
* {{cite web |last=Bartlett |first=Troy |date=18 December 2017 |url=https://bugguide.net/node/view/194 |title=Family Lycaenidae - Blues, Coppers, Hairstreaks, Harvesters |website=BugGuide |accessdate=5 February 2020}}
* {{cite web |last=Bartlett |first=Troy |date=18 December 2017 |url=https://bugguide.net/node/view/194 |title=Family Lycaenidae Blues, Coppers, Hairstreaks, Harvesters |website=BugGuide |access-date=5 February 2020}}
* [http://insecta.pro/family/lycaenidae/ "Family Lycaenidae"]. ''Insecta.pro''.
* [https://insecta.pro/taxonomy/92 "Family Lycaenidae"]. ''Insecta.pro''.
* {{NCBI|124406}}
* {{NCBI taxid|124406}}
* {{ITIS|ID=694072|''Lycaeninae'' [Leach], 1815}}
* {{ITIS|ID=694072|''Lycaeninae'' [Leach], 1815}}
* {{Eol|2682619}}
* {{Eol|2682619}}
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[[Category:Butterfly families]]
[[Category:Butterfly families]]
[[Category:Taxa named by William Elford Leach]]
[[Category:Taxa named by William Elford Leach]]
[[Category:Papilionoidea]]

Latest revision as of 13:02, 8 March 2024

Lycaenidae
The large blue, a lycaenid butterfly
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Superfamily: Papilionoidea
Family: Lycaenidae
Leach, 1815
Subfamilies

and see text or List of lycaenid genera

Brown hairstreak (Thecla betulae)
Loxura atymnus

Lycaenidae is the second-largest family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide,[1] whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfly species.

The family comprises seven subfamilies, including the blues (Polyommatinae), the coppers (Lycaeninae), the hairstreaks (Theclinae), and the harvesters (Miletinae).

Description, food, and life cycle[edit]

Adults are small, under 5 cm usually, and brightly coloured, sometimes with a metallic gloss. Lycaenidae wings are generally blue or green. More than half of these butterflies depend on ants in some way.[2]

Larvae are often flattened rather than cylindrical, with glands that may produce secretions that attract and subdue ants. Their cuticles tend to be thickened. Some larvae are capable of producing vibrations and low sounds that are transmitted through the substrates they inhabit. They use these sounds to communicate with ants.[3][4]

Adult individuals often have hairy antenna-like tails complete with black and white annulated (ringed) appearance. Many species also have a spot at the base of the tail and some turn around upon landing to confuse potential predators from recognizing the true head orientation. This causes predators to approach from the true head end resulting in early visual detection or to attack the false head ending up with a beak of dusty scales.[5]

Lycaenid caterpillars are diverse in their food habits and apart from phytophagy, some are entomophagous, feeding on aphids, scale insects, and ant larvae. Some lycaenids even exploit their association with ants by inducing ants to feed them by regurgitation, a process called trophallaxis. Not all lycaenid butterflies need ants, but about 75% of species associate with ants,[3] a relationship called myrmecophily. These associations can be mutualistic, parasitic, or predatory depending on the species.

In some species, larvae are attended and protected by ants while feeding on the host plant, and the ants receive sugar-rich honeydew from them, throughout the larval life, and in some species during the pupal stage. In other species, only the first few instars are spent on the plant, and the remainder of the larval lifespan is spent as a predator within the ant nest. It becomes a parasite, feeding on ant regurgitations, or a predator on the ant larvae.[3] The caterpillars pupate inside the ants' nest and the ants continue to look after the pupae. Just before the adults emerge, the wings of the butterfly inside the pupal case detach from it, and the pupa becomes silvery. The adult butterfly emerges from the pupa after three to four weeks, still inside the ant nest. The butterfly must crawl out of the ant nest before it can expand its wings.

Several evolutionary adaptations enable these associations, including small glands on the skin of the caterpillars called "pore cupola organs". Caterpillars of many species have a gland on the seventh abdominal segment that produces honeydew and is called the "dorsal nectary gland" (also called "Newcomer's gland"). An eversible organ called the "tentacular organ" is present on the eighth abdominal segment and this is cylindrical and topped with a ring of spikes and emits chemical signals which are believed to help in communicating with ants.[6]

Subfamilies[edit]

Mating Cyaniris semiargus

Many taxonomists only include the Lycaeninae, Theclinae, Polyommatinae, Poritiinae, Miletinae, and Curetinae under the Lycaenidae.[7][8] The Aphnaeinae, which used to be a tribe (Aphnaeini) within the Theclinae, were recently given subfamily rank too.[9]

Some older classifications used to include other subfamilies such as Liphyrinae (now Liphyrini, a tribe within Miletinae), Lipteninae (now Liptenini, a tribe within Poritiinae), or Riodininae (now a separate family: Riodinidae).

The fossil genus Lithodryas is usually (but not unequivocally) placed here; Lithopsyche is sometimes placed here, but sometimes in the Riodininae.

See also[edit]

References[edit]

  1. ^ Pierce, Naomi E.; Braby, Michael F.; Heath, Alan; Lohman, David J.; Mathew, John; Rand, Douglas B.; Travassos, Mark A. (January 2002). "The Ecology and Evolution of Ant Association in the Lycaenidae (Lepidoptera)". Annual Review of Entomology. 47 (1): 733–771. doi:10.1146/annurev.ento.47.091201.145257. PMID 11729090.
  2. ^ Ueda, Shouhei; Komatsu, Takashi; Itino, Takao; Arai, Ryusuke; Sakamoto, Hironori (3 November 2016). "Host-ant specificity of endangered large blue butterflies (Phengaris spp., Lepidoptera: Lycaenidae) in Japan". Scientific Reports. 6 (1): 36364. Bibcode:2016NatSR...636364U. doi:10.1038/srep36364. PMC 5093462. PMID 27808223.
  3. ^ a b c Pierce, Naomi E.; Braby, Michael F.; Heath, Alan; Lohman, David J.; Mathew, John; Rand, Douglas B.; Travassos, Mark A. (January 2002). "The Ecology and Evolution of Ant Association in the Lycaenidae (Lepidoptera)". Annual Review of Entomology. 47 (1): 733–771. doi:10.1146/annurev.ento.47.091201.145257. PMID 11729090. ProQuest 222345887.
  4. ^ DeVries, Philip J. (1992). "Singing Caterpillars, Ants and Symbiosis". Scientific American. 267 (4): 76–83. Bibcode:1992SciAm.267d..76D. doi:10.1038/scientificamerican1092-76. JSTOR 24939256.
  5. ^ Robbins, Robert K. (1981). "The 'False Head' Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies". The American Naturalist. 118 (5): 770–775. doi:10.1086/283868. JSTOR 2460614. S2CID 34146954.
  6. ^ "Lycaenid Butterflies and Ants". Australian Museum. Archived 18 November 2007.
  7. ^ Brower, Andrew V. Z. (2008). "Lycaenidae [Leach] 1815". Version 25 April 2008 (under construction). The Tree of Life Web Project.
  8. ^ Ackery, P. R.; de Jong, R. & Vane-Wright, R. I. (1999). "The butterflies: Hedyloidea, Hesperioidea, and Papilionoidea". Pages 264-300 in: Lepidoptera: Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. Handbook of Zoology Vol. IV, Part 35. N. P. Kristensen, ed. De Gruyter, Berlin and New York.
  9. ^ Boyle, John H.; Kaliszewska, Zofia A.; Espeland, Marianne; Suderman, Tamara R.; Fleming, Jake; Heath, Alan; Pierce, Naomi E. (January 2015). "Phylogeny of the A phnaeinae: myrmecophilous A frican butterflies with carnivorous and herbivorous life histories". Systematic Entomology. 40 (1): 169–182. Bibcode:2015SysEn..40..169B. doi:10.1111/syen.12098. S2CID 83052979.

Further reading[edit]

External links[edit]