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{{Short description|Action of animals deliberately ejecting blood from their bodies}}
'''Autohaemorrhaging''' or '''reflex bleeding''' is the action of insects deliberately ejecting [[haemolymph]] from themselves. If the insect has toxic compounds in their blood then it may be an effective chemical defense. Insects that autohaemorrhage include:<ref name="autohaem">{{cite journal|last=Bateman|first=Philip|coauthors=P. A. Fleming|date=28 Apr 2009|title=There will be blood: autohaemorrhage behaviour as part of the defence repertoire of an insect|journal=Journal of Zoology|volume=278|issue=4|pages=342–348|issn=1469-7998|url=http://www3.interscience.wiley.com/journal/122368498/abstract|accessdate=07-08-2009}}</ref>
[[File:Phrynosoma orbiculare.JPG|thumb|[[Horned lizard]] showing evidence of autohaemorrhaging]]
'''Autohaemorrhaging,''' or '''reflex bleeding,''' is the action of animals deliberately ejecting [[blood]] from their bodies. Autohaemorrhaging has been observed as occurring in two variations.<ref name=":0">{{Cite journal|last1=Iftime|first1=Alexandru|last2=Iftime|first2=Oana|date=January 2014|title=Thanatosis and autohaemorrhaging in the Aesculapian Snake Zamenis|url=https://www.zobodat.at/publikation_articles.php?id=200376|journal=Herpetozoa|volume=26|issue=3–4|pages=173–174|via=Zobodat}}</ref> In the first form, blood is squirted toward a predator. The blood of these animals usually contains toxic compounds, making the behaviour an effective chemical [[anti-predator adaptation|defense mechanism]]. In the second form, blood is not squirted, but is slowly emitted from the animal's body. This form appears to serve a deterrent effect, and is used by animals whose blood does not seem to be toxic.<ref name=":0" /> Most animals that autohaemorrhage are insects, but some reptiles also display this behaviour.<ref name=":03">{{Cite journal|last1=Bateman|first1=P. W.|last2=Fleming|first2=P. A.|date=2009|title=There will be blood: autohaemorrhage behaviour as part of the defence repertoire of an insect|journal=Journal of Zoology|language=en|volume=278|issue=4|pages=342–348|doi=10.1111/j.1469-7998.2009.00582.x|issn=1469-7998}}</ref>


Some organisms have shown an ability to tailor their autohaemorrhaging response. Armoured crickets will projectile autohaemorrhage over longer distances when attacked from the side, compared to being attacked from an overhead predator.<ref name=":1">{{Cite journal|last1=Bateman|first1=Philip W.|last2=Fleming|first2=Patricia A.|date=2012-04-26|title=Signaling or Not-Signaling: Variation in Vulnerability and Defense Tactics of Armored Ground Crickets (Acanthoplus Speiseri: Orthoptera, Tettigoniidae, Hetrodinae)|journal=Journal of Insect Behavior|volume=26|issue=1|pages=14–22|doi=10.1007/s10905-012-9329-5|s2cid=18599346|issn=0892-7553}}</ref>
==Beetles==
*[[Blister beetle]]s
*[[Meloidae]] whose haemolymph contains [[cantharidin]] that they sequester from plants that they feed on.
*''[[Timarcha]]'' species
⚫
*[[Chrysomelidae]] whose haemolymph contains [[anthraquinone]]s.
*[[Ladybug]]s<ref>[http://books.google.co.uk/books?id=yoK7APpgbqoC&pg=PA202&lpg=PA202&dq=Autohaemorrhage+ladybug&source=bl&ots=Edko9i08Tp&sig=MeTDsErbvEDriLXLXhIC0-aKcSg&hl=en&ei=NZnLSpLdHsyH4gabxOyIBg&sa=X&oi=book_result&ct=result&resnum=1#v=onepage&q=&f=false The Alkaloids: chemistry and physiology, Volume 31 By Arnold Brossi]</ref>


==Flies==
== Insects ==
Several orders of insects have been observed to utilize this defence mechanism.
*[[Sawfly]] larvae
*[[Stonefly]] larvae


==[[Orthoptera]]==
*[[Beetle]]s
**[[Meloidae]] (blister beetles) – their haemolymph contains [[cantharidin]] that they sequester from plants on which they feed. One of the known species is ''[[Meloe americanus]]''.<ref name="Seasonal Activity of Meloe americanus">{{cite journal |last1=Young |first1=O.P. |title=Seasonal Activity of Meloe americanus (Coleoptera: Meloidae) in a Mississippi Old-field Habitat |journal=Journal of Entomological Science |date=October 1995 |volume=30 |issue=4 |pages=434–442 |doi=10.18474/0749-8004-30.4.434 |url=https://doi.org/10.18474/0749-8004-30.4.434}}</ref>
⚫
*[[Bushhopper]]s such as ''[[Dictyophora spumans]]'' and ''[[Phymateus leprosus]]'' whose haemolymph contains cardiac [[glycosides]] sequestered from [[milkweed]] on which they feed.
**[[Tenebrionidae]] (darkling beetles) - larvae of ''[[Asbolus verrucosus]]'' have been observed to autohaemorrhage while they feign death.<ref>{{cite journal |last1=Rider |first1=S. Dean |last2=Hostetler |first2=Heather A. |title=Reflex Bleeding in Tonically Immobilized Larvae Causes Debris-Based Camouflage in the Blue Death-Feigning Beetle, Asbolus verrucosus LeConte (Coleoptera: Tenebrionidae) |journal=The Coleopterists Bulletin |date=21 June 2022 |volume=76 |issue=2 |page=237-247 |doi=10.1649/0010-065X-76.2.237|s2cid=249480941 }}</ref>
⚫
*[[Katydid]]s including ''[[Eugaster]]'' species
⚫
**[[Chrysomelidae]], incl. ''[[Timarcha]]'' species - their haemolymph contains [[anthraquinone]]s.
*[[Armoured ground cricket]]s
**[[Coccinellidae]] (ladybird, ladybug or lady beetles) – An alkaloid toxin in the haemolymph is exuded through the joints of the [[exoskeleton]], triggered by mechanical stimulation (such as predator attack).<ref>[https://books.google.com/books?id=yoK7APpgbqoC&dq=Autohaemorrhage+ladybug&pg=PA202 The Alkaloids: chemistry and physiology, Volume 31 By Arnold Brossi]</ref>
*''[[Enyaliopsis nyala]]''
*[[Hemiptera]]
**[[Cercopidae]]<ref>{{cite journal |last1=Peck |first1=Daniel C. |title=Reflex Bleeding in Froghoppers (Homoptera: Cercopidae): Variation in Behavior and Taxonomic Distribution |journal=Annals of the Entomological Society of America |date=1 September 2000 |volume=93 |issue=5 |pages=1186–1194 |doi=10.1603/0013-8746(2000)093[1186:RBIFHC]2.0.CO;2 |url=https://academic.oup.com/aesa/article/93/5/1186/9145?login=true}}</ref> – including [[spittlebugs]] ''[[Prosapia bicincta]]'' and ''Prosapia ignipectus''<ref>{{Cite journal |doi = 10.1155/2016/3623092|title = Abrupt Geographical Transition between Aposematic Color Forms in the Spittlebug ''Prosapia'' ignipectus(Fitch) (Hemiptera: Cercopidae)|journal = Psyche: A Journal of Entomology|volume = 2016|pages = 1–10|year = 2016|last1 = Thompson|first1 = Vinton|last2 = Carvalho|first2 = Gervasio S.|doi-access = free}}</ref>
*[[Lepidoptera]]
**[[Arctiinae|Tiger moth]] adults, such as ''[[Arctia caja]]'', which mixes haemolymph with glandular products (neurotoxic choline esters).<ref>V.H. Resh, R.T. Cardé (2003) ''Encyclopedia of Insects''. p. 168</ref>
*[[Orthoptera]]
⚫
**[[Bushhopper]]s, such as ''[[Dictyophorus spumans]]'', ''[[Phymateus viridipes]]'' and ''[[Phymateus leprosus]]'' – their haemolymph contains cardiac [[glycosides]], sequestered from [[milkweed]] on which they feed.
⚫
**[[Katydid]]s, including ''[[Eugaster]]'' species
**[[Armoured ground cricket]]s<ref>{{cite journal |last1=van Huis |first1=Arnold |title=Cultural significance of locusts, grasshoppers, and crickets in sub-Saharan Africa |journal=Journal of Ethnobiology and Ethnomedicine |date=26 March 2022 |volume=18 |issue=1 |pages=24 |doi=10.1186/s13002-022-00524-w |doi-access=free |pmid=35346258 |pmc=8962220 |language=en |issn=1746-4269}}</ref>
*[[Plecoptera]]
**[[Stonefly]] larvae <ref>{{cite journal |last1=Moore |first1=Kathryn A. |last2=Williams |first2=D. Dudley |title=Novel Strategies in the Complex Defense Repertoire of a Stonefly (Pteronarcys dorsata) nymph |journal=Oikos |date=1990 |volume=57 |issue=1 |pages=49–56 |doi=10.2307/3565735 |jstor=3565735 |bibcode=1990Oikos..57...49M |hdl=1807/762 |url=https://www.jstor.org/stable/3565735 |issn=0030-1299|hdl-access=free }}</ref>

== Reptiles ==
[[File:Tropidophis_curtus_barbouri_autohaemorrhaging.jpg|thumb|A West Indian wood snake displaying autohaemorrhaging. The eyes are fully flooded with blood and some is emerging from the mouth.]]

=== Lizards ===

*[[Horned lizard|Horned lizards]] ([[Phrynosomatidae]]). At least six species of horned lizards are able to squirt an aimed stream of blood from the corners of their eyes, up to 5 feet (1.5 m).<ref>{{Cite journal|last1=Sherbrooke|first1=Wade C.|last2=Middendorf III|first2=George A.|date=2001|title=Blood-Squirting Variability in Horned Lizards (Phrynosoma)|url=https://bioone.org/journals/copeia/volume-2001/issue-4/0045-8511(2001)001[1114:BSVIHL]2.0.CO;2/Blood-Squirting-Variability-in-Horned-Lizards-Phrynosoma/10.1643/0045-8511(2001)001[1114:BSVIHL]2.0.CO;2.full|journal=Copeia|volume=2001|issue=4|pages=1114–1122|doi=10.1643/0045-8511(2001)001[1114:BSVIHL]2.0.CO;2|s2cid=86061329 |issn=0045-8511}}</ref>

=== Snakes ===

* West Indian wood snake (''[[Tropidophis]]''). Thirteen species have been found to expel blood from the mouth and nostrils while also fully flooding both eyes with blood.<ref>{{Cite journal|last1=Hoefer|first1=Sebastian|last2=Mills|first2=Sophie|last3=Robinson|first3=Nathan J.|date=2019|title=Autohaemorrhaging in a Bahamian pygmy boa, Tropidophis curtus barbouri|url=https://www.thebhs.org/publications/the-herpetological-bulletin/issue-number-150-winter-2019/2018-13-autohaemorrhaging-in-a-bahamian-pygmy-boa-i-tropidophis-curtus-barbouri-i|journal= Herpetological Bulletin|language=en-gb|issue=150|pages=39–40|doi=10.33256/hb150.3940|doi-access=free}}</ref>
* European grass snake (''[[Natrix natrix]]''), which secretes blood from the lining of the mouth while playing dead.<ref>{{Cite journal|last1=Gregory|first1=Patrick T.|last2=Isaac|first2=Leigh Anne|last3=Griffiths|first3=Richard A|date=2007|title=Death feigning by grass snakes (Natrix natrix) in response to handling by human "predators."|url=http://doi.apa.org/getdoi.cfm?doi=10.1037/0735-7036.121.2.123|journal=Journal of Comparative Psychology|language=en|volume=121|issue=2|pages=123–129|doi=10.1037/0735-7036.121.2.123|pmid=17516791|issn=1939-2087}}</ref>
* Long-nosed snake (''[[Long-nosed snake|Rhinocheilus lecontei]]''), which exudes blood from the cloaca.<ref name=":2">{{Cite journal|last1=Smith|first1=Donald D.|last2=Pflanz|first2=Deborah J.|last3=Powell|first3=Robert|date=1993|title=Observations of autohemorrhaging in Tropidophis haetianus, Rhinocheilus lecontei, Heterodon platyrhinos, and Nerodia erythrogaster|journal=Herpetological Review|volume=24|pages=130–131}}</ref>
* Eastern hognose snake ([[Eastern hognose snake|''Heterodon'' ''platirhinos'']]), which emits blood from the cloacal region.<ref name=":2" />
* Plain-bellied water snake (''[[Nerodia erythrogaster]]''), which releases blood from the mouth.<ref name=":2" />
* Western hognose snake (''[[Heterodon nasicus]]''), which releases blood from the mouth.

== Consequences of reflexive bleeding ==
In some cases, the loss of blood can be substantial. Beetles may lose up to 13% of their net body weight as a consequence of expelling haemolymph.<ref>{{Cite book|last=Klowden|first=Marc J.|date=October 2007|publisher=Elsevier|volume=2|pages=369|doi=10.1016/c2011-0-04120-0|isbn=9780124158191|title=Physiological Systems in Insects|s2cid=63715931 }}</ref> Autohaemorrhaging may result in dehydration. The ejection of blood puts organisms at risk of cannibalism from other members of their species.<ref name=":12">{{Cite journal|last1=Bateman|first1=Philip W.|last2=Fleming|first2=Patricia A.|date=2012-04-26|title=Signaling or Not-Signaling: Variation in Vulnerability and Defense Tactics of Armored Ground Crickets (Acanthoplus Speiseri: Orthoptera, Tettigoniidae, Hetrodinae)|journal=Journal of Insect Behavior|volume=26|issue=1|pages=14–22|doi=10.1007/s10905-012-9329-5|s2cid=18599346|issn=0892-7553}}</ref>
[[File:Canibalism - Reflex bleeding (2106637702).jpg|thumb|An inactive [[prepupa]] [[Harmonia axyridis|Asian ladybeetle]] autohaemorrhaging, resulting from cannibalism by an adult of the same species.]]


==See also==
==See also==
⚫
*[[Haemorrhage]]
*[[Autothysis]]
*[[Autothysis]]
⚫
*[[Haemorrhage]]


==References==
==References==
{{reflist}}
{{reflist}}
== External links ==
*{{commonscat-inline|Autohaemorrhaging|Autohaemorrhaging}}


[[Category:Antipredator adaptations]]
[[Category:Blood]]
[[Category:Blood]]
[[Category:Entomology]]
[[Category:Insect physiology]]

Latest revision as of 12:24, 22 November 2024

Horned lizard showing evidence of autohaemorrhaging

Autohaemorrhaging, or reflex bleeding, is the action of animals deliberately ejecting blood from their bodies. Autohaemorrhaging has been observed as occurring in two variations.[1] In the first form, blood is squirted toward a predator. The blood of these animals usually contains toxic compounds, making the behaviour an effective chemical defense mechanism. In the second form, blood is not squirted, but is slowly emitted from the animal's body. This form appears to serve a deterrent effect, and is used by animals whose blood does not seem to be toxic.[1] Most animals that autohaemorrhage are insects, but some reptiles also display this behaviour.[2]

Some organisms have shown an ability to tailor their autohaemorrhaging response. Armoured crickets will projectile autohaemorrhage over longer distances when attacked from the side, compared to being attacked from an overhead predator.[3]

Insects

[edit]

Several orders of insects have been observed to utilize this defence mechanism.

Reptiles

[edit]
A West Indian wood snake displaying autohaemorrhaging. The eyes are fully flooded with blood and some is emerging from the mouth.

Lizards

[edit]
  • Horned lizards (Phrynosomatidae). At least six species of horned lizards are able to squirt an aimed stream of blood from the corners of their eyes, up to 5 feet (1.5 m).[12]

Snakes

[edit]
  • West Indian wood snake (Tropidophis). Thirteen species have been found to expel blood from the mouth and nostrils while also fully flooding both eyes with blood.[13]
  • European grass snake (Natrix natrix), which secretes blood from the lining of the mouth while playing dead.[14]
  • Long-nosed snake (Rhinocheilus lecontei), which exudes blood from the cloaca.[15]
  • Eastern hognose snake (Heterodon platirhinos), which emits blood from the cloacal region.[15]
  • Plain-bellied water snake (Nerodia erythrogaster), which releases blood from the mouth.[15]
  • Western hognose snake (Heterodon nasicus), which releases blood from the mouth.

Consequences of reflexive bleeding

[edit]

In some cases, the loss of blood can be substantial. Beetles may lose up to 13% of their net body weight as a consequence of expelling haemolymph.[16] Autohaemorrhaging may result in dehydration. The ejection of blood puts organisms at risk of cannibalism from other members of their species.[17]

An inactive prepupa Asian ladybeetle autohaemorrhaging, resulting from cannibalism by an adult of the same species.

See also

[edit]

References

[edit]
  1. ^ a b Iftime, Alexandru; Iftime, Oana (January 2014). "Thanatosis and autohaemorrhaging in the Aesculapian Snake Zamenis". Herpetozoa. 26 (3–4): 173–174 – via Zobodat.
  2. ^ Bateman, P. W.; Fleming, P. A. (2009). "There will be blood: autohaemorrhage behaviour as part of the defence repertoire of an insect". Journal of Zoology. 278 (4): 342–348. doi:10.1111/j.1469-7998.2009.00582.x. ISSN 1469-7998.
  3. ^ Bateman, Philip W.; Fleming, Patricia A. (2012-04-26). "Signaling or Not-Signaling: Variation in Vulnerability and Defense Tactics of Armored Ground Crickets (Acanthoplus Speiseri: Orthoptera, Tettigoniidae, Hetrodinae)". Journal of Insect Behavior. 26 (1): 14–22. doi:10.1007/s10905-012-9329-5. ISSN 0892-7553. S2CID 18599346.
  4. ^ Young, O.P. (October 1995). "Seasonal Activity of Meloe americanus (Coleoptera: Meloidae) in a Mississippi Old-field Habitat". Journal of Entomological Science. 30 (4): 434–442. doi:10.18474/0749-8004-30.4.434.
  5. ^ Rider, S. Dean; Hostetler, Heather A. (21 June 2022). "Reflex Bleeding in Tonically Immobilized Larvae Causes Debris-Based Camouflage in the Blue Death-Feigning Beetle, Asbolus verrucosus LeConte (Coleoptera: Tenebrionidae)". The Coleopterists Bulletin. 76 (2): 237-247. doi:10.1649/0010-065X-76.2.237. S2CID 249480941.
  6. ^ The Alkaloids: chemistry and physiology, Volume 31 By Arnold Brossi
  7. ^ Peck, Daniel C. (1 September 2000). "Reflex Bleeding in Froghoppers (Homoptera: Cercopidae): Variation in Behavior and Taxonomic Distribution". Annals of the Entomological Society of America. 93 (5): 1186–1194. doi:10.1603/0013-8746(2000)093[1186:RBIFHC]2.0.CO;2.
  8. ^ Thompson, Vinton; Carvalho, Gervasio S. (2016). "Abrupt Geographical Transition between Aposematic Color Forms in the Spittlebug Prosapia ignipectus(Fitch) (Hemiptera: Cercopidae)". Psyche: A Journal of Entomology. 2016: 1–10. doi:10.1155/2016/3623092.
  9. ^ V.H. Resh, R.T. Cardé (2003) Encyclopedia of Insects. p. 168
  10. ^ van Huis, Arnold (26 March 2022). "Cultural significance of locusts, grasshoppers, and crickets in sub-Saharan Africa". Journal of Ethnobiology and Ethnomedicine. 18 (1): 24. doi:10.1186/s13002-022-00524-w. ISSN 1746-4269. PMC 8962220. PMID 35346258.
  11. ^ Moore, Kathryn A.; Williams, D. Dudley (1990). "Novel Strategies in the Complex Defense Repertoire of a Stonefly (Pteronarcys dorsata) nymph". Oikos. 57 (1): 49–56. Bibcode:1990Oikos..57...49M. doi:10.2307/3565735. hdl:1807/762. ISSN 0030-1299. JSTOR 3565735.
  12. ^ Sherbrooke, Wade C.; Middendorf III, George A. (2001). "Blood-Squirting Variability in Horned Lizards (Phrynosoma)". Copeia. 2001 (4): 1114–1122. doi:10.1643/0045-8511(2001)001[1114:BSVIHL]2.0.CO;2. ISSN 0045-8511. S2CID 86061329.
  13. ^ Hoefer, Sebastian; Mills, Sophie; Robinson, Nathan J. (2019). "Autohaemorrhaging in a Bahamian pygmy boa, Tropidophis curtus barbouri". Herpetological Bulletin (150): 39–40. doi:10.33256/hb150.3940.
  14. ^ Gregory, Patrick T.; Isaac, Leigh Anne; Griffiths, Richard A (2007). "Death feigning by grass snakes (Natrix natrix) in response to handling by human "predators."". Journal of Comparative Psychology. 121 (2): 123–129. doi:10.1037/0735-7036.121.2.123. ISSN 1939-2087. PMID 17516791.
  15. ^ a b c Smith, Donald D.; Pflanz, Deborah J.; Powell, Robert (1993). "Observations of autohemorrhaging in Tropidophis haetianus, Rhinocheilus lecontei, Heterodon platyrhinos, and Nerodia erythrogaster". Herpetological Review. 24: 130–131.
  16. ^ Klowden, Marc J. (October 2007). Physiological Systems in Insects. Vol. 2. Elsevier. p. 369. doi:10.1016/c2011-0-04120-0. ISBN 9780124158191. S2CID 63715931.
  17. ^ Bateman, Philip W.; Fleming, Patricia A. (2012-04-26). "Signaling or Not-Signaling: Variation in Vulnerability and Defense Tactics of Armored Ground Crickets (Acanthoplus Speiseri: Orthoptera, Tettigoniidae, Hetrodinae)". Journal of Insect Behavior. 26 (1): 14–22. doi:10.1007/s10905-012-9329-5. ISSN 0892-7553. S2CID 18599346.
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