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{{Short description|Family of insects}}
{{Automatic taxobox
{{Automatic taxobox
| taxon = Boreidae
| taxon = Boreidae
| fossil_range = {{fossilrange|Tithonian|Present|[[Late Jurassic]] to [[Present]]}}
| fossil_range =
|authority=| image = Boreus hyemalis female.jpg
| authority =
| image = Boreus hyemalis female.jpg
| image_caption = ''Boreus hiemalis''
| image_caption = ''[[Boreus hyemalis]]''
| subdivision_ranks = Genera
| subdivision_ranks = Genera
| subdivision =
| subdivision = * ''[[Boreus]]''
*''[[Boreus]]''
* ''[[Caurinus]]''
*''[[Caurinus]]''
* ''[[Hesperoboreus]]''
*''[[Hesperoboreus]]''
}}
}}


'''Boreidae''', commonly called '''snow scorpionflies''', or in the [[British Isles]], '''snow fleas''' (no relation to the snow flea ''[[Hypogastrura nivicola]]'') are a very small family of [[Scorpionfly|scorpionflies]], containing only around 30 species, all of which are [[boreal ecosystem|boreal]] or high-altitude species in the [[Northern Hemisphere]]. Recent research indicates the boreids are more closely related to [[flea]]s than to other scorpionflies, which renders the order [[Mecoptera]] [[paraphyletic]] if the order [[Siphonaptera]] is excluded from it.<ref name="Fleas">{{cite journal|
'''Boreidae''', commonly called '''snow scorpionflies''', or in the [[British Isles]], '''snow fleas''' (no relation to the snow flea ''[[Hypogastrura nivicola]]'') are a very small family of [[Scorpionfly|scorpionflies]], containing only around 30 species, all of which are [[boreal ecosystem|boreal]] or high-altitude species in the [[Northern Hemisphere]].
author=Whiting, M. F.|title=Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera |year = 2002 |
journal = Zoologica Scripta |volume = 31 |issue = 1 |pages = 93–104 |doi=10.1046/j.0300-3256.2001.00095.x|url=https://semanticscholar.org/paper/d3537d8be461ca0419bc02920d0f4d69b50c0214 }}</ref>


These insects are small (typically 6&nbsp;mm or less), with the wings reduced to bristles or absent, and they are somewhat compressed, so in fact some resemblance to fleas is noted. They are most commonly active during the winter months, towards the transition into spring, and the larvae and adults typically feed on mosses. The adults will often disperse between breeding areas by walking across the open snow, thus the common name. The males use their bristle-like wings to help grasp the female over his back while mating, while the wings of females are vestigial small oval pads with no ability to allow her to fly. The adults have a long [[Rostrum (anatomy)|rostrum]] formed from the [[clypeus (arthropod anatomy)|clypeus]] and [[labrum (arthropod mouthpart)|labrum]], [[genae]], and [[maxillo-labium]].
These [[insect]]s are small (typically 6&nbsp;mm or less), with the wings reduced to bristles or absent, and they are somewhat compressed, so in fact some resemblance to fleas is noted. They are most commonly active during the winter months, towards the transition into spring, and the larvae and adults typically feed on mosses. The adults will often disperse between breeding areas by walking across the open snow, thus the common name. The males use their bristle-like wings to help grasp the female over their backs while mating, while the wings of females are vestigial small oval pads with no ability to allow them to fly. The adults have a long [[Rostrum (anatomy)|rostrum]] formed from the [[clypeus (arthropod anatomy)|clypeus]] and [[labrum (arthropod mouthpart)|labrum]], [[genae]], and [[Maxilla (arthropod mouthpart)|maxillo]]-[[Labium (insect mouthpart)|labium]].


The body temperature, and therefore activity level, of this scorpionfly depends on its absorption of short-wave and long-wave radiation rather than surrounding air temperatures (by which it is completely unaffected). The boundary layer of snow that the insect occupies has very low thermal conductance, and so the insect loses its own heat very slowly here. This delicate balance between cold and heat means that the animal is easily killed by heat when held in a human hand.<ref>{{cite book|author=Daniel Marlos|title=The Curious World of Bugs: The Bugman's Guide to the Mysterious and Remarkable Lives of Things That Crawl|url=https://books.google.com/books?id=icyVPVBVnUQC&pg=PT218|date=2010-10-05|publisher=Penguin Publishing Group|isbn=978-1-101-44406-1|page=218}}</ref>
The body temperature, and therefore activity level, of this scorpionfly depends on its absorption of short-wave and long-wave radiation rather than surrounding air temperatures (by which it is completely unaffected). The boundary layer of snow that the insect occupies has very low thermal conductance, and so the insect loses its own heat very slowly here. This delicate balance between cold and heat means that the animal is easily killed by heat when held in a human hand.<ref>{{cite book|author=Daniel Marlos|title=The Curious World of Bugs: The Bugman's Guide to the Mysterious and Remarkable Lives of Things That Crawl|url=https://books.google.com/books?id=icyVPVBVnUQC&pg=PT218|date=2010-10-05|publisher=Penguin Publishing Group|isbn=978-1-101-44406-1|page=218}}</ref>

The group has been proposed in some studies to be the closest relatives of [[fleas]] (Siphonaptera), rendering Mecoptera paraphyletic. This has been disputed by other studies, which find [[Nannochoristidae]] more closely related to fleas instead.<ref name=":0">{{Cite journal |last1=Tihelka |first1=Erik |last2=Giacomelli |first2=Mattia |last3=Huang |first3=Di-Ying |last4=Pisani |first4=Davide |last5=Donoghue |first5=Philip C. J. |last6=Cai |first6=Chen-Yang |date=2020-12-21 |title=Fleas are parasitic scorpionflies |url=https://www.mapress.com/pe/article/view/palaeoentomology.3.6.16 |journal=Palaeoentomology |language=en |volume=3 |issue=6 |pages=641–653–641–653 |doi=10.11646/palaeoentomology.3.6.16 |issn=2624-2834 |hdl-access=free |hdl=1983/8d3c12c6-529c-4754-b59d-3abf88a32fc9 |s2cid=234423213}}</ref>


== Phylogeny ==
== Phylogeny ==


It is unclear as of 2020 whether the Mecoptera form a single clade, or whether the [[Siphonaptera]] (fleas) are inside that clade, so that the traditional "Mecoptera" is paraphyletic. However the earlier suggestion that the Siphonaptera are sister to the Boreidae<ref name=Whiting2008>{{cite journal |last1=Whiting |first1=Michael F. |author1-link=Michael F. Whiting|last2=Whiting |first2=Alison S. |last3=Hastriter |first3=Michael W. |last4=Dittmar |first4=Katharina |title=A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations |journal=Cladistics |date=2008 |volume=24 |issue=5 |pages=677–707 |doi=10.1111/j.1096-0031.2008.00211.x |citeseerx=10.1.1.731.5211 |s2cid=33808144 }}</ref><ref name=Whiting2002>{{cite journal |last1=Whiting |first1=Michael F. |author1-link=Michael F. Whiting|year=2002 |title=Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera |journal=Zoologica Scripta |volume=31 |issue=1 |pages=93–104 |url=http://www3.interscience.wiley.com/journal/118916123/abstract |archive-url=https://archive.today/20130105095659/http://www3.interscience.wiley.com/journal/118916123/abstract |url-status=dead |archive-date=2013-01-05 |doi=10.1046/j.0300-3256.2001.00095.x |s2cid=56100681 }}</ref><ref name=WiegmannYates2012>{{cite book |last1=Wiegmann |first1=Brian |last2=Yeates |first2=David K. |title=The Evolutionary Biology of Flies |url=https://books.google.com/books?id=rElP5sNn6IYC&pg=PA5 |year=2012 |publisher=Columbia University Press |isbn=978-0-231-50170-5 |page=5 |quote=Recently, a close affinity between Siphonaptera and Mecoptera has been convincingly demonstrated via morphology (Bilinski et al. 1998) and molecular data (Whiting 2002), rendering Mecoptera paraphyletic, but making the clade including Mecoptera and Siphonaptera monophyletic}}</ref> is not supported. The two possible trees are shown below:<ref name="Meusemann Trautwein 2020">{{cite journal |last1=Meusemann |first1=Karen |last2=Trautwein |first2=Michelle |last3=Friedrich |first3=Frank |last4=Beutel |first4=Rolf G. |last5=Wiegmann |first5=Brian M. |last6=Donath |first6=Alexander |last7=Podsiadlowski |first7=Lars |last8=Petersen |first8=Malte |last9=Niehuis |first9=Oliver |last10=Mayer |first10=Christoph |last11=Bayless |first11=Keith M. |last12=Shin |first12=Seunggwan |last13=Liu |first13=Shanlin |last14=Hlinka |first14=Ondrej |last15=Minh |first15=Bui Quang |last16=Kozlov |first16=Alexey |last17=Morel |first17=Benoit |last18=Peters |first18=Ralph S. |last19=Bartel |first19=Daniela |last20=Grove |first20=Simon |last21=Zhou |first21=Xin |last22=Misof |first22=Bernhard |last23=Yeates |first23=David K. |title=Are Fleas Highly Modified Mecoptera? Phylogenomic Resolution of Antliophora (Insecta: Holometabola) |journal=Bioarchiv |year=2020 |doi=10.1101/2020.11.19.390666}}</ref>
It is unclear as of 2020 whether the Mecoptera form a single clade, or whether the [[Siphonaptera]] (fleas) are inside that clade, so that the traditional "Mecoptera" is paraphyletic. However the earlier suggestion that the Siphonaptera are sister to the Boreidae<ref name=Whiting2008>{{cite journal |last1=Whiting |first1=Michael F. |author1-link=Michael F. Whiting|last2=Whiting |first2=Alison S. |last3=Hastriter |first3=Michael W. |last4=Dittmar |first4=Katharina |title=A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations |journal=Cladistics |date=2008 |volume=24 |issue=5 |pages=677–707 |doi=10.1111/j.1096-0031.2008.00211.x |citeseerx=10.1.1.731.5211 |s2cid=33808144 }}</ref><ref name=Whiting2002>{{cite journal |last1=Whiting |first1=Michael F. |author1-link=Michael F. Whiting|year=2002 |title=Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera |journal=Zoologica Scripta |volume=31 |issue=1 |pages=93–104 |url=http://www3.interscience.wiley.com/journal/118916123/abstract |archive-url=https://archive.today/20130105095659/http://www3.interscience.wiley.com/journal/118916123/abstract |url-status=dead |archive-date=2013-01-05 |doi=10.1046/j.0300-3256.2001.00095.x |s2cid=56100681 }}</ref><ref name=WiegmannYates2012>{{cite book |last1=Wiegmann |first1=Brian |last2=Yeates |first2=David K. |title=The Evolutionary Biology of Flies |url=https://books.google.com/books?id=rElP5sNn6IYC&pg=PA5 |year=2012 |publisher=Columbia University Press |isbn=978-0-231-50170-5 |page=5 |quote=Recently, a close affinity between Siphonaptera and Mecoptera has been convincingly demonstrated via morphology (Bilinski et al. 1998) and molecular data (Whiting 2002), rendering Mecoptera paraphyletic, but making the clade including Mecoptera and Siphonaptera monophyletic}}</ref> is not supported. The two possible trees are shown below:<ref name="Meusemann Trautwein 2020">{{cite bioRxiv |last1=Meusemann |first1=Karen |last2=Trautwein |first2=Michelle |last3=Friedrich |first3=Frank |last4=Beutel |first4=Rolf G. |last5=Wiegmann |first5=Brian M. |last6=Donath |first6=Alexander |last7=Podsiadlowski |first7=Lars |last8=Petersen |first8=Malte |last9=Niehuis |first9=Oliver |last10=Mayer |first10=Christoph |last11=Bayless |first11=Keith M. |last12=Shin |first12=Seunggwan |last13=Liu |first13=Shanlin |last14=Hlinka |first14=Ondrej |last15=Minh |first15=Bui Quang |last16=Kozlov |first16=Alexey |last17=Morel |first17=Benoit |last18=Peters |first18=Ralph S. |last19=Bartel |first19=Daniela |last20=Grove |first20=Simon |last21=Zhou |first21=Xin |last22=Misof |first22=Bernhard |last23=Yeates |first23=David K. |display-authors=5|title=Are Fleas Highly Modified Mecoptera? Phylogenomic Resolution of Antliophora (Insecta: Holometabola) |year=2020 |biorxiv=10.1101/2020.11.19.390666}}</ref>


(a) Mecoptera is paraphyletic, Boriedae is sister to ([[Nannochoristidae]] + [[Siphonaptera]]):<ref name="Meusemann Trautwein 2020"/>
(a) Mecoptera is paraphyletic, Boreidae is sister to ([[Nannochoristidae]] + [[Siphonaptera]]):<ref name="Meusemann Trautwein 2020" />


{{clade
{{clade
|label1=[[Antliophora]]
|label1=[[Antliophora]]
|1={{clade
|1={{clade
|1=[[Diptera]] (true flies) [[File:Common house fly, Musca domestica.jpg|70px]]
|1=[[Diptera]] (true flies) [[File:Common house fly, Musca domestica.jpg|70px]]
|2={{clade
|2={{clade
|1=[[Pistillifera]] (scorpionflies, hangingflies, 400 [[Species|spp]].) [[File:Gunzesrieder Tal Insekt 3.jpg|90px]]
|1=[[Pistillifera]] (scorpionflies, hangingflies, 400 [[Species|spp]].) [[File:Gunzesrieder Tal Insekt 3.jpg|90px]]
Line 42: Line 43:
}}
}}


(b) Mecoptera is monophyletic, Boreidae is sister to [[Pistillifera]]:<ref name="Meusemann Trautwein 2020"/>
(b) Mecoptera is monophyletic, Boreidae is sister to [[Pistillifera]]:<ref name="Meusemann Trautwein 2020" />


{{clade
{{clade
|label1=[[Antliophora]]
|label1=[[Antliophora]]
|1={{clade
|1={{clade
|label1=[[Mecoptera]]
|label1=[[Mecoptera]]
|1={{clade
|1={{clade
Line 55: Line 56:
|2=[[Nannochoristidae]] (southern scorpionflies, 8 spp.)
|2=[[Nannochoristidae]] (southern scorpionflies, 8 spp.)
}}
}}
|label2=[[Siphonaptera]]
|label2=[[Siphonaptera]]
|2= (fleas, 2500 spp.) [[File:Pulex irritans female ZSM.jpg|50px]]
|2= (fleas, 2500 spp.) [[File:Pulex irritans female ZSM.jpg|50px]]
}}
}}
Line 62: Line 63:
== Genera ==
== Genera ==
This list is adapted from the ''World Checklist of extant Mecoptera species'',<ref name="Cat1997">[http://www.calacademy.org/research/entomology/Entomology_Resources/mecoptera/boreidae.htm Boreidae] {{webarchive|url=https://archive.today/20040111035237/http://www.calacademy.org/research/entomology/Entomology_Resources/mecoptera/boreidae.htm |date=2004-01-11 }}</ref> and is complete as of 1997. The number of species in each genus is indicated in parentheses.
This list is adapted from the ''World Checklist of extant Mecoptera species'',<ref name="Cat1997">[http://www.calacademy.org/research/entomology/Entomology_Resources/mecoptera/boreidae.htm Boreidae] {{webarchive|url=https://archive.today/20040111035237/http://www.calacademy.org/research/entomology/Entomology_Resources/mecoptera/boreidae.htm |date=2004-01-11 }}</ref> and is complete as of 1997. The number of species in each genus is indicated in parentheses.

* ''[[Boreus]]'' (24) <small>Latreille, 1816</small> (North America, Europe, Asia)
* ''[[Boreus]]'' (24) <small>Latreille, 1816</small> (North America, Europe, Asia)
** ''[[Boreus hyemalis]]'' - also called the snow flea.
** ''[[Boreus hyemalis]]'' also called the snow flea.
* ''[[Caurinus]]'' (2) <small>Russell, 1979</small> (Oregon, Alaska)<ref>{{cite journal|last=Sikes|first=Derek|author2=Jill Stockbridge |title=Description of Caurinus tlagu, new species, from Prince of Wales Island, Alaska (Mecoptera, Boreidae, Caurininae)|journal=ZooKeys|date=July 11, 2013|issue=316|pages=35–53|doi=10.3897/zookeys.316.5400|url=https://www.pensoft.net/journals/zookeys/article/5400/description-of-caurinus-tlagu-new-species-from-prince-of-wales-island-alaska-mecoptera-boreidae-caurininae-|pmid=23878513|pmc=3713333}}</ref>
* ''[[Caurinus]]'' (2) <small>Russell, 1979</small> (Oregon, Alaska)<ref>{{cite journal|last=Sikes|first=Derek|author2=Jill Stockbridge |title=Description of Caurinus tlagu, new species, from Prince of Wales Island, Alaska (Mecoptera, Boreidae, Caurininae)|journal=ZooKeys|date=July 11, 2013|issue=316|pages=35–53|doi=10.3897/zookeys.316.5400|url=https://www.pensoft.net/journals/zookeys/article/5400/description-of-caurinus-tlagu-new-species-from-prince-of-wales-island-alaska-mecoptera-boreidae-caurininae-|pmid=23878513|pmc=3713333|doi-access=free}}</ref>
* ''[[Hesperoboreus]]'' (2) <small>Penny, 1977</small> (USA)
* ''[[Hesperoboreus]]'' (2) <small>Penny, 1977</small> (United States)


== See also ==
== See also ==
* [[Glacier flea]]
* [[Glacier flea]]
* [[Snow flies]] genus ''Chionea'' - a [[Convergent evolution|convergent]] genus of wingless [[crane fly|crane flies]]
* [[Snow flies]] genus ''Chionea'' a [[Convergent evolution|convergent]] genus of wingless [[crane fly|crane flies]]
* [[Apteropanorpidae]] - another family of wingless scorpionflies
* [[Apteropanorpidae]] another family of wingless scorpionflies


== References ==
== References ==
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{{Taxonbar|from=Q2004861}}
{{Taxonbar|from=Q2004861}}


[[Category:Snow scorpionflies| ]]
[[Category:Mecoptera]]
[[Category:Mecoptera]]

Latest revision as of 23:53, 12 December 2023

Snow scorpionfly
Boreus hyemalis
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Mecoptera
Family: Boreidae
Genera

Boreidae, commonly called snow scorpionflies, or in the British Isles, snow fleas (no relation to the snow flea Hypogastrura nivicola) are a very small family of scorpionflies, containing only around 30 species, all of which are boreal or high-altitude species in the Northern Hemisphere.

These insects are small (typically 6 mm or less), with the wings reduced to bristles or absent, and they are somewhat compressed, so in fact some resemblance to fleas is noted. They are most commonly active during the winter months, towards the transition into spring, and the larvae and adults typically feed on mosses. The adults will often disperse between breeding areas by walking across the open snow, thus the common name. The males use their bristle-like wings to help grasp the female over their backs while mating, while the wings of females are vestigial small oval pads with no ability to allow them to fly. The adults have a long rostrum formed from the clypeus and labrum, genae, and maxillo-labium.

The body temperature, and therefore activity level, of this scorpionfly depends on its absorption of short-wave and long-wave radiation rather than surrounding air temperatures (by which it is completely unaffected). The boundary layer of snow that the insect occupies has very low thermal conductance, and so the insect loses its own heat very slowly here. This delicate balance between cold and heat means that the animal is easily killed by heat when held in a human hand.[1]

The group has been proposed in some studies to be the closest relatives of fleas (Siphonaptera), rendering Mecoptera paraphyletic. This has been disputed by other studies, which find Nannochoristidae more closely related to fleas instead.[2]

Phylogeny[edit]

It is unclear as of 2020 whether the Mecoptera form a single clade, or whether the Siphonaptera (fleas) are inside that clade, so that the traditional "Mecoptera" is paraphyletic. However the earlier suggestion that the Siphonaptera are sister to the Boreidae[3][4][5] is not supported. The two possible trees are shown below:[6]

(a) Mecoptera is paraphyletic, Boreidae is sister to (Nannochoristidae + Siphonaptera):[6]

Antliophora

Diptera (true flies)

Pistillifera (scorpionflies, hangingflies, 400 spp.)

Nannochoristidae (southern scorpionflies, 8 spp.)

Siphonaptera (fleas, 2500 spp.)

Boreidae (snow scorpionflies, 30 spp.)

(b) Mecoptera is monophyletic, Boreidae is sister to Pistillifera:[6]

Antliophora
Mecoptera

Pistillifera (scorpionflies, hangingflies, 400 spp.)

Boreidae (snow scorpionflies, 30 spp.)

Nannochoristidae (southern scorpionflies, 8 spp.)

Siphonaptera

(fleas, 2500 spp.)

Genera[edit]

This list is adapted from the World Checklist of extant Mecoptera species,[7] and is complete as of 1997. The number of species in each genus is indicated in parentheses.

See also[edit]

References[edit]

  1. ^ Daniel Marlos (2010-10-05). The Curious World of Bugs: The Bugman's Guide to the Mysterious and Remarkable Lives of Things That Crawl. Penguin Publishing Group. p. 218. ISBN 978-1-101-44406-1.
  2. ^ Tihelka, Erik; Giacomelli, Mattia; Huang, Di-Ying; Pisani, Davide; Donoghue, Philip C. J.; Cai, Chen-Yang (2020-12-21). "Fleas are parasitic scorpionflies". Palaeoentomology. 3 (6): 641–653–641–653. doi:10.11646/palaeoentomology.3.6.16. hdl:1983/8d3c12c6-529c-4754-b59d-3abf88a32fc9. ISSN 2624-2834. S2CID 234423213.
  3. ^ Whiting, Michael F.; Whiting, Alison S.; Hastriter, Michael W.; Dittmar, Katharina (2008). "A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations". Cladistics. 24 (5): 677–707. CiteSeerX 10.1.1.731.5211. doi:10.1111/j.1096-0031.2008.00211.x. S2CID 33808144.
  4. ^ Whiting, Michael F. (2002). "Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera". Zoologica Scripta. 31 (1): 93–104. doi:10.1046/j.0300-3256.2001.00095.x. S2CID 56100681. Archived from the original on 2013-01-05.
  5. ^ Wiegmann, Brian; Yeates, David K. (2012). The Evolutionary Biology of Flies. Columbia University Press. p. 5. ISBN 978-0-231-50170-5. Recently, a close affinity between Siphonaptera and Mecoptera has been convincingly demonstrated via morphology (Bilinski et al. 1998) and molecular data (Whiting 2002), rendering Mecoptera paraphyletic, but making the clade including Mecoptera and Siphonaptera monophyletic
  6. ^ a b c Meusemann, Karen; Trautwein, Michelle; Friedrich, Frank; Beutel, Rolf G.; Wiegmann, Brian M.; et al. (2020). "Are Fleas Highly Modified Mecoptera? Phylogenomic Resolution of Antliophora (Insecta: Holometabola)". bioRxiv 10.1101/2020.11.19.390666.
  7. ^ Boreidae Archived 2004-01-11 at archive.today
  8. ^ Sikes, Derek; Jill Stockbridge (July 11, 2013). "Description of Caurinus tlagu, new species, from Prince of Wales Island, Alaska (Mecoptera, Boreidae, Caurininae)". ZooKeys (316): 35–53. doi:10.3897/zookeys.316.5400. PMC 3713333. PMID 23878513.