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== Black-Headed Gull ==
The black-headed gull displays a variety of compelling behaviours and adaptations. Some of these include removing eggshells from ones nest after hatching, begging coordination between siblings, differences between sexes, conspecific brood parasitism, and extra-pair paternity.

== Behaviour ==
== Behaviour ==


=== Eggshell Removal ===
=== Eggshell Removal ===
Eggshell removal is a behaviour seen in birds once the chicks have hatched to reduce risk of predation<ref name=":0" />. Removing the eggshell acts as a way of camouflage to avoid predators seeing the nest<ref name=":1">{{Cite journal|last=Houghton|first=J.C.W.|last2=Feekes|first2=F.|last3=Broekhuysen|first3=G.J.|last4=Tinbergen|first4=N.|last5=Szulc|first5=E.|last6=Kruuk|first6=H.|date=1962|title=Egg Shell Removal By the Black-Headed Gull, Larus Ridibundus L.; a Behaviour Component of Camouflage|url=http://dx.doi.org/10.1163/156853961x00213|journal=Behaviour|volume=19|issue=1-2|pages=74–116|doi=10.1163/156853961x00213|issn=0005-7959}}</ref>. The predation risk while egg shells are present decreases the further away they are from the nest (once removed)<ref name=":0">{{Cite journal|last=SORDAHL|first=TEX A.|date=2006|title=FIELD EXPERIMENTS ON EGGSHELL REMOVAL BY MOUNTAIN PLOVERS|url=http://dx.doi.org/10.1676/1559-4491(2006)118[0059:feoerb]2.0.co;2|journal=The Wilson Journal of Ornithology|volume=118|issue=1|pages=59–63|doi=10.1676/1559-4491(2006)118[0059:feoerb]2.0.co;2|issn=1559-4491|via=}}</ref>. Black-headed gull eggs experience predation from different species of birds, foxes, stoats, and even other black-headed gulls. Although mothers show some form of aggressiveness when a predator is near, wet chicks can be easily taken by other black-headed gulls after hatching when the parents of the wet chick are distracted<ref name=":1" />.
Eggshell removal is a behaviour seen in birds once the chicks have hatched to reduce risk of predation<ref name=":0" />. Removing the eggshell acts as a way of camouflage to avoid predators seeing the nest<ref name=":1">{{Cite journal|last=Houghton|first=J.C.W.|last2=Feekes|first2=F.|last3=Broekhuysen|first3=G.J.|last4=Tinbergen|first4=N.|last5=Szulc|first5=E.|last6=Kruuk|first6=H.|date=1962|title=Egg Shell Removal By the Black-Headed Gull, Larus Ridibundus L.; a Behaviour Component of Camouflage|url=http://dx.doi.org/10.1163/156853961x00213|journal=Behaviour|volume=19|issue=1-2|pages=74–116|doi=10.1163/156853961x00213|issn=0005-7959}}</ref>. The predation risk while egg shells are present decreases the further away they are from the nest (once removed)<ref name=":0">{{Cite journal|last=SORDAHL|first=TEX A.|date=2006|title=FIELD EXPERIMENTS ON EGGSHELL REMOVAL BY MOUNTAIN PLOVERS|url=http://dx.doi.org/10.1676/1559-4491(2006)118[0059:feoerb]2.0.co;2|journal=The Wilson Journal of Ornithology|volume=118|issue=1|pages=59–63|doi=10.1676/1559-4491(2006)118[0059:feoerb]2.0.co;2|issn=1559-4491|via=}}</ref>. Black-headed gull eggs experience predation from different species of birds, foxes, stoats, and even other black-headed gulls. Although mothers show some form of aggressiveness when a predator is near, in the first 30 minutes, wet chicks can be easily taken by other black-headed gulls after hatching when the parents of the wet chick are distracted<ref name=":1" />.


Black headed gulls also carry away other objects that do not belong in the nest. The removal and eggshells and other objects is important not only in the incubation period but also during the few days when the eggs hatch. However, the removal has the tendency to increase as time goes on<ref>{{Cite journal|last=Beer|first=C.G.|date=1963|title=Incubation and Nest-Building Behaviour of Black-Headed Gulls Iv: Nest-Building in the Laying and Incubation Periods|url=http://dx.doi.org/10.1163/156853963x00158|journal=Behaviour|volume=21|issue=3-4|pages=155–176|doi=10.1163/156853963x00158|issn=0005-7959}}</ref>. The removal is done by both the male and female parents, normally lasts a few seconds and is done three times a year<ref name=":1" />.
Black headed gulls also carry away other objects that do not belong in the nest. The removal of eggshells and other objects is important not only in the incubation period but also during the first few days after the eggs hatch. However, the removal process seems to increase as time goes on<ref>{{Cite journal|last=Beer|first=C.G.|date=1963|title=Incubation and Nest-Building Behaviour of Black-Headed Gulls Iv: Nest-Building in the Laying and Incubation Periods|url=http://dx.doi.org/10.1163/156853963x00158|journal=Behaviour|volume=21|issue=3-4|pages=155–176|doi=10.1163/156853963x00158|issn=0005-7959}}</ref>. The removal is done by both the male and female parents, normally lasts a few seconds and is done three times a year<ref name=":1" />.


==== Earlier Eggshell Removal Hypotheses ====
==== Earlier Eggshell Removal Hypotheses ====
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==== Conspecific Brood Parasitism ====
==== Conspecific Brood Parasitism ====
Conspecific brood parasitism is a behaviour where females lay their eggs in another females nest, of the same species. It can reduce the cost of incubation and nestling young by passing it on to another bird. Black-headed gulls usually lay three egg clutches, and the first two are normally larger than the third. The third egg normally has the lowest survival rate, while the first or second are usually the parasitic eggs <ref name=":3">{{Cite journal|last=Duda|first=Norbert|last2=Chętnicki|first2=Włodzimierz|date=2012|title=Conspecific Brood Parasitism is Biased Towards Relatives in the Common Black-Headed Gull|url=http://dx.doi.org/10.5253/078.100.0110|journal=Ardea|volume=100|issue=1|pages=63–70|doi=10.5253/078.100.0110|issn=0373-2266|via=}}</ref>.
Conspecific brood parasitism is a behaviour that occurs when females lay their eggs in another females nest, of the same species<ref name=":3" />. It can reduce the cost of incubation and nestling young by passing it on to another bird. Black-headed gulls usually lay three egg clutches, and the first two are normally larger than the third. The third egg normally has the lowest survival rate, while the first or second are usually the parasitic eggs <ref name=":3">{{Cite journal|last=Duda|first=Norbert|last2=Chętnicki|first2=Włodzimierz|date=2012|title=Conspecific Brood Parasitism is Biased Towards Relatives in the Common Black-Headed Gull|url=http://dx.doi.org/10.5253/078.100.0110|journal=Ardea|volume=100|issue=1|pages=63–70|doi=10.5253/078.100.0110|issn=0373-2266|via=}}</ref>.


Most of the egg dumping occurs within the beginning of the egg laying period. The parasitic eggs being laid in another conspecifics nest increases the chance of hatching and may occur because of nest desertion or a nest being taken over by another bird<ref name=":3" />.
Most of the egg dumping occurs within the beginning of the egg laying period. The parasitic eggs being laid in another conspecifics nest increases the chance of hatching and may occur because of nest desertion or a nest being taken over by another bird<ref name=":3" />.
Line 37: Line 40:


==== Extra-Pair Paternity ====
==== Extra-Pair Paternity ====
The rate of EPP has a large variation between populations of black-headed gulls. In black-headed gulls, extra-pair paternity is primarily a context-dependent strategy, meaning not all birds of this species experience this behaviour<ref>{{Cite journal|last=Indykiewicz|first=Piotr|last2=Podlaszczuk|first2=Patrycja|last3=Minias|first3=Piotr|date=2017|title=Extra-pair paternity in the black-headed gull: is it exceptional among colonial waterbirds?|url=http://dx.doi.org/10.1163/1568539x-00003459|journal=Behaviour|volume=154|issue=11|pages=1081–1099|doi=10.1163/1568539x-00003459|issn=0005-7959}}</ref>. The variation between populations of extra-pair paternity can be explained by the variation it has on the advantages and disadvantages it has on a female, as well, as the variation in constraints on a females choice <ref>{{Cite journal|last=Petrie|first=Marion|last2=Kempenaers|first2=Bart|date=1998|title=Extra-pair paternity in birds: explaining variation between species and populations|url=http://dx.doi.org/10.1016/s0169-5347(97)01232-9|journal=Trends in Ecology & Evolution|volume=13|issue=2|pages=52–58|doi=10.1016/s0169-5347(97)01232-9|issn=0169-5347|via=}}</ref>.
The rate of extra-pair paternity (EPP) has a large variation between populations of black-headed gulls. It is primarily a context-dependent strategy, meaning not all birds of this species experience this behaviour<ref>{{Cite journal|last=Indykiewicz|first=Piotr|last2=Podlaszczuk|first2=Patrycja|last3=Minias|first3=Piotr|date=2017|title=Extra-pair paternity in the black-headed gull: is it exceptional among colonial waterbirds?|url=http://dx.doi.org/10.1163/1568539x-00003459|journal=Behaviour|volume=154|issue=11|pages=1081–1099|doi=10.1163/1568539x-00003459|issn=0005-7959}}</ref>. The variation between populations of extra-pair paternity can be explained by the variation it has on the advantages and disadvantages it has on a female, as well, as the variation in pressure on a females choice <ref>{{Cite journal|last=Petrie|first=Marion|last2=Kempenaers|first2=Bart|date=1998|title=Extra-pair paternity in birds: explaining variation between species and populations|url=http://dx.doi.org/10.1016/s0169-5347(97)01232-9|journal=Trends in Ecology & Evolution|volume=13|issue=2|pages=52–58|doi=10.1016/s0169-5347(97)01232-9|issn=0169-5347|via=}}</ref>.


Nests have been seen to carry eggs from more than one mother at a time. This may occur as a result of intraspecifc nest parasitism<ref name=":4" />, joint female nesting<ref name=":5" /> and nest takeover<ref name=":6" />. Intraspecific nest parasitism is a cost to hosts because the female could end up taking care of the parasitic chicks over her own and therefore neglecting them and reducing their fitness. Another disadvantage for the host is that incubating more chicks than their own takes up more energy<ref name=":7" />.
Nests have been seen to carry eggs from more than one mother at a time. This may occur as a result of intra-specific nest parasitism<ref name=":4" />, joint female nesting<ref name=":5" /> and nest takeover<ref name=":6" />. Intra-specific nest parasitism is a cost to hosts because the female could end up taking care of the parasitic chicks over her own and therefore neglecting them and reducing their fitness. Another disadvantage for the host is that incubating more chicks than their own takes up more energy<ref name=":7" />.


== References ==
== References ==

Revision as of 15:37, 15 November 2019

Black-Headed Gull

The black-headed gull displays a variety of compelling behaviours and adaptations. Some of these include removing eggshells from ones nest after hatching, begging coordination between siblings, differences between sexes, conspecific brood parasitism, and extra-pair paternity.

Behaviour

Eggshell Removal

Eggshell removal is a behaviour seen in birds once the chicks have hatched to reduce risk of predation[1]. Removing the eggshell acts as a way of camouflage to avoid predators seeing the nest[2]. The predation risk while egg shells are present decreases the further away they are from the nest (once removed)[1]. Black-headed gull eggs experience predation from different species of birds, foxes, stoats, and even other black-headed gulls. Although mothers show some form of aggressiveness when a predator is near, in the first 30 minutes, wet chicks can be easily taken by other black-headed gulls after hatching when the parents of the wet chick are distracted[2].

Black headed gulls also carry away other objects that do not belong in the nest. The removal of eggshells and other objects is important not only in the incubation period but also during the first few days after the eggs hatch. However, the removal process seems to increase as time goes on[3]. The removal is done by both the male and female parents, normally lasts a few seconds and is done three times a year[2].

Earlier Eggshell Removal Hypotheses

Earlier hypotheses have attempted to explain the survival value of black-headed gulls removing their eggshells from the nest, including[2]:

  1. The sharp edges of the shells after hatching could harm the chicks
  2. The eggshell could somehow intrude during the brooding
  3. The eggshell could slip over the unhatched egg, creating a double shell
  4. Some of the moist organic material left from the shell could lead to a production of bacteria and mould

Walking Displays

Black-headed gulls display both head-bobbing walking (HBW) and non-bobbing walking (NBW). Head-bobbing walking is expressed by a hold phase and a thrust phase. The hold phase in black-headed gulls occurs mainly during the single support phase and is when the bird balances its head to equal the environment. Head-bobbing walking occurs during seeking type foraging by wading and includes benefits such as enhancing motion and pattern detection and gathering depth information from motion parallax during the thrust phase. Non-bobbing walking occurs when black-headed gulls are displaying a waiting behaviour while foraging on flat substrate[4].

Breeding

Begging Coordination Between Siblings

Black-headed gulls feed their young by regurgitating it onto the ground, rather than into each chick one at a time. The parents tend to accommodate their regurgitation amounts for how intense the nest begging is, from both an individual chick or a group of chicks begging together. Chicks who are siblings, have learned this behaviour and begin synchronizing their begging signals to limit individual effort [5]. The rate of parental food regurgitation to chicks increases with begging intensity[6].

The amount and response of begging signals differs throughout the nestling period. Usually, there is 3-5 begging events/hour, each lasting around 1 minute. High intensity begging behaviour appears at the end of the first week in the nest, but the coordination between multiple chicks emerge during the last week of the nestling period. The more siblings present, the more they coordinate their begging while decreasing the number of begging[5].

Sex Differences

Male chicks have less of a chance of survival when compared to female chicks. Black-headed gulls are a sexually size-dimorphic species, so the larger sex is at a disadvantage when the amount of food sources are low[7].

Male birds are more likely to be born in the first egg and female birds are more likely to be born in the third. The position of a female black-headed gull in response to food availability when laying the eggs can predict the offspring future prospects[8].

Conspecific Brood Parasitism

Conspecific brood parasitism is a behaviour that occurs when females lay their eggs in another females nest, of the same species[9]. It can reduce the cost of incubation and nestling young by passing it on to another bird. Black-headed gulls usually lay three egg clutches, and the first two are normally larger than the third. The third egg normally has the lowest survival rate, while the first or second are usually the parasitic eggs [9].

Most of the egg dumping occurs within the beginning of the egg laying period. The parasitic eggs being laid in another conspecifics nest increases the chance of hatching and may occur because of nest desertion or a nest being taken over by another bird[9].

Multiple eggs in a nest from different mothers may also result from intraspecifc nest parasitism[10], joint female nesting[11], and nest takeover[12]. Intraspecific nest parasitism is a disadvantage to the hosts because the female could end up taking care of the parasitic chicks over her own and therefore neglecting them and reducing their fitness. Another disadvantage for the host is that incubating more chicks than their own takes up more energy[13].

Extra-Pair Paternity

The rate of extra-pair paternity (EPP) has a large variation between populations of black-headed gulls. It is primarily a context-dependent strategy, meaning not all birds of this species experience this behaviour[14]. The variation between populations of extra-pair paternity can be explained by the variation it has on the advantages and disadvantages it has on a female, as well, as the variation in pressure on a females choice [15].

Nests have been seen to carry eggs from more than one mother at a time. This may occur as a result of intra-specific nest parasitism[10], joint female nesting[11] and nest takeover[12]. Intra-specific nest parasitism is a cost to hosts because the female could end up taking care of the parasitic chicks over her own and therefore neglecting them and reducing their fitness. Another disadvantage for the host is that incubating more chicks than their own takes up more energy[13].

References

  1. ^ a b SORDAHL, TEX A. (2006). "FIELD EXPERIMENTS ON EGGSHELL REMOVAL BY MOUNTAIN PLOVERS". The Wilson Journal of Ornithology. 118 (1): 59–63. doi:10.1676/1559-4491(2006)118[0059:feoerb]2.0.co;2. ISSN 1559-4491.
  2. ^ a b c d Houghton, J.C.W.; Feekes, F.; Broekhuysen, G.J.; Tinbergen, N.; Szulc, E.; Kruuk, H. (1962). "Egg Shell Removal By the Black-Headed Gull, Larus Ridibundus L.; a Behaviour Component of Camouflage". Behaviour. 19 (1–2): 74–116. doi:10.1163/156853961x00213. ISSN 0005-7959.
  3. ^ Beer, C.G. (1963). "Incubation and Nest-Building Behaviour of Black-Headed Gulls Iv: Nest-Building in the Laying and Incubation Periods". Behaviour. 21 (3–4): 155–176. doi:10.1163/156853963x00158. ISSN 0005-7959.
  4. ^ Fujita, Masaki (2006-01-24). "Head-bobbing and non-bobbing walking of black-headed gulls (Larus ridibundus)". Journal of Comparative Physiology A. 192 (5): 481–488. doi:10.1007/s00359-005-0083-4. ISSN 0340-7594.
  5. ^ a b Blanc, Alain; Ogier, Nicolas; Roux, Angélique; Denizeau, Sébastien; Mathevon, Nicolas (2010). "Begging coordination between siblings in Black-headed Gulls". Comptes Rendus Biologies. 333 (9): 688–693. doi:10.1016/j.crvi.2010.06.002. ISSN 1631-0691.
  6. ^ Mathevon, N.; Charrier, I. (2004-05-07). "Parent–offspring conflict and the coordination of siblings in gulls". Proceedings of the Royal Society of London. Series B: Biological Sciences. 271 (suppl_4). doi:10.1098/rsbl.2003.0117. ISSN 0962-8452.
  7. ^ MULLER, WENDT; GROOTHUIS, TON G. G.; EISING, CORINE M.; DIJKSTRA, COR (2005). "An experimental study on the causes of sex-biased mortality in the black-headed gull - the possible role of testosterone". Journal of Animal Ecology. 74 (4): 735–741. doi:10.1111/j.1365-2656.2005.00964.x. ISSN 0021-8790.
  8. ^ Lezalova, Radka; Tkadlec, Emil; Obornik, Miroslav; Simek, Jaroslav; Honza, Marcel (2005-09-07). "Should males come first? The relationship between offspring hatching order and sex in the black-headed gull Larus ridibundus". Journal of Avian Biology. 0 (0): 060118052425010–???. doi:10.1111/j.2005.0908-8857.03466.x. ISSN 0908-8857.
  9. ^ a b c Duda, Norbert; Chętnicki, Włodzimierz (2012). "Conspecific Brood Parasitism is Biased Towards Relatives in the Common Black-Headed Gull". Ardea. 100 (1): 63–70. doi:10.5253/078.100.0110. ISSN 0373-2266.
  10. ^ a b YOM-TOV, YORAM (2008-06-28). "An updated list and some comments on the occurrence of intraspecific nest parasitism in birds". Ibis. 143 (1): 133–143. doi:10.1111/j.1474-919x.2001.tb04177.x. ISSN 0019-1019.
  11. ^ a b Vehrencamp, Sandra L. (2000). "Evolutionary routes to joint-female nesting in birds". Behavioral Ecology. 11 (3): 334–344. doi:10.1093/beheco/11.3.334. ISSN 1465-7279.
  12. ^ a b Waldeck, Peter; Andersson, Malte (2006). "Brood Parasitism and Nest Takeover in Common Eiders". Ethology. 112 (6): 616–624. doi:10.1111/j.1439-0310.2005.01187.x. ISSN 0179-1613.
  13. ^ a b Duda, Norbert; Ch e ˛ tnicki, Wlodzimierz; Waldeck, Peter; Andersson, Malte (2008-01-07). "Multiple maternity in black-headed gull Larus ridibundus clutches as revealed by protein fingerprinting". Journal of Avian Biology. 0 (0): 080205233540538–0. doi:10.1111/j.2007.0908-8857.04111.x. ISSN 0908-8857.
  14. ^ Indykiewicz, Piotr; Podlaszczuk, Patrycja; Minias, Piotr (2017). "Extra-pair paternity in the black-headed gull: is it exceptional among colonial waterbirds?". Behaviour. 154 (11): 1081–1099. doi:10.1163/1568539x-00003459. ISSN 0005-7959. {{cite journal}}: no-break space character in |title= at position 50 (help)
  15. ^ Petrie, Marion; Kempenaers, Bart (1998). "Extra-pair paternity in birds: explaining variation between species and populations". Trends in Ecology & Evolution. 13 (2): 52–58. doi:10.1016/s0169-5347(97)01232-9. ISSN 0169-5347.
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