Interacciones intraespecíficas durante la alimentación en grupo en la Gaviota de Olrog (Larus atlanticus)

Francisco Zumpano; Germán Oscar García

doi:10.56178/eh.v37i1.361

Articles Published 01 August 2022 https://doi.org/10.56178/eh.v37i1.361

Intra-specific interactions during group feeding in the Olrog’s Gull (Larus atlanticus)

Argentina foraging kleptoparasitism Larus atlanticus seabirds trophic displacements

Francisco Zumpano Instituto de Investigaciones Marinas y Costeras (IIMyC), Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata – CONICET, Rodríguez Peña 4046 Nivel 1, B7602GSD Mar del Plata, Argentina

Germán Oscar García Instituto de Investigaciones Marinas y Costeras (IIMyC), Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata – CONICET, Rodríguez Peña 4046 Nivel 1, B7602GSD Mar del Plata, Argentina

Abstract

In this study, we analyzed the intra-specific interactions that occur between individuals of Olrog’s Gull (Larus atlanticus) while foraging in groups outside the breeding season. Interactions were studied by analyzing behavioral observations of foraging individuals (n=171) at the Mar Chiquita Coastal Lagoon (Buenos Aires Province, Argentina). We recorded a total of 98 trophic displacements events and 70 kleptoparasitic events, in which successful attacks were 95% and 24% respectively. Juveniles and males initiated de interactions in the 77% of the trophic displacements and 65% of the kleptoparasitic events. The attacks were “with physical contact” in the 55% and 32% of the trophic displacements and kleptoparasitic events, respectively. The main tactic response for both intra-specific interactions was “avoiding”. The probability of occurring a kleptoparasitism event was related to the prey size. We conclude that juveniles and males were the main initiators of the interactions studied and that the main difference between them was the success rate. This study is the first to analyze and discuss the intra-specific interactions that occur between individuals of Olrog’s Gull. Understanding their feeding behavior is important to discover new aspects of natural history and to provide background information for a threatened endemic species.

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References

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aMat ja (1990) Food usurpation by waterfowl and wa-ders. Wildfowl 41:107-116
annorbah nnd y holbech lh (2012) Relative abundan- ce, agonistic behaviour, and resource partitioning among three scavenging bird species in Ghana. Ma- limbus 34:1-8
azPiroz ab, alFaro M y jiMénez s (2012) Lista Roja delas Aves del Uruguay. Una evaluación del estado de conservación de la avifauna nacional con base en los criterios de la Unión Internacional para la Con- servación de la Naturaleza. Dirección Nacional de Medio Ambiente, Montevideo
barnard cj (1984) The evolution of food-scroungingstrategies within and between species. Pp. 95-126 en: barnard cj (ed) Producers and Scroungers: Stra- tegies of Exploitation and Parasitism. Croom Helm, Londres
bates d, Mächler M, bolKer b y walKer s (2015) FittingLinear Mixed-Effects Models Using lme4. Journal ofStatistical Software 67(1): 1-48
bautista lM, alonso jc y alonso ja (1998) Foraging site displacement in common crane flocks. Animal Be- haviour 56:1237-1243
beauchaMP G (2017) Disentangling the various mecha-nisms that account for the decline in vigilance with group size. Behavioural Processes 136:59-63
beauchaMP G (2019) On how risk and group size in-teract to influence vigilance. Biological Reviews94:1918-1934
berón MP, García Go, luPPi t y Favero M (2011) Age-re- lated prey selectivity and foraging efficiency of Olrog’s Gulls (Larus atlanticus) feeding on crabs in their non-breeding grounds. Emu 111:172-178
bertellotti nM y yorio P (2000) Age-related feedingbehaviour and foraging efficiency in Kelp Gulls La- rus dominicanus attending coastal trawlers in Argen- tina. Ardea 88(2): 207-214
birdliFe international (2021) Species factsheet: Larus atlanticus (URL http://www.birdlife.org)
boschi ee (1964) Los crustáceos decápodos Brachyu- ra del litoral bonaerense (R. Argentina). Boletín del Instituto de Biología Marina 6:1-99
bower KM (2003). When to use Fisher’s exact test.
American Society for Quality, Six Sigma Forum Magazi-ne 2(4): 35-37.
brocKMann hj y barnard cj (1979) Kleptoparasitism in birds. Animal behaviour 27:487-514
burGer j (1988) Foraging behavior in gulls: differen-ces in method, prey, and habitat. Colonial Waterbirds9-23.
burGer j y GochFeld M (1979) Age differences in Ring-billed Gull kleptoparasitism on Starlings. The Auk 96(4): 806-808.
castano M, biondi lM, zuMPano F, Favero M y García Go
(2020) Behavioral responses to a novel feeding pro- blem in the Olrog’s Gull Larus atlanticus. Marine Or- nithology 48:149-155
coPello s y Favero M (2001) Foraging ecology of Olrog’sGull Larus atlanticus in Mar Chiquita Lagoon (Bue- nos Aires, Argentina): are there age-related diffe- rences? Bird Conservation International 11:175-188
crawley Mj (2007) The R Book. John Wiley and Sons,UK.
duriez o, herMan s y sarrazin F (2012) Intra-specific competition in foraging Griffon Vultures Gyps ful- vus: 2. The influence of supplementary feeding ma- nagement. Bird Study 59:193-206
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Favero M (1996) Foraging ecology of Pale-Faced Shea-thbills in colonies of southern Elephant Seals at King George Island, Antarctica. Journal of Field Or- nithology 67:292-299
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Flower tP y Gribble M (2012) Kleptoparasitism by at-tacks versus false alarm calls in fork-tailed dron- gos. Animal Behaviour 83:403-410
Flower tP, child MF y ridley ar (2013) The ecologicaleconomics of kleptoparasitism: pay‐offs from self‐ foraging versus kleptoparasitism. Journal of Animal Ecology 82:245-255
Furness rw (1987) Kleptoparasitism in seabirds. Pp.
77-99 en: croxall jP (ed) Seabirds: Feeding Biology and Role in Marine Ecosystems. Cambridge Univer- sity Press, Cambridge
GaGlio d, sherley rb, cooK tr, ryan PG y Flower t(2018) The costs of kleptoparasitism: a study of mixed-species seabird breeding colonies. Behavio- ral Ecology 29:939-947
García Go y biondi lM (2011) Kleptoparasitism by theCaracara Chimango (Milvago chimango) on the Ame- rican Oystercatcher (Haematopus palliatus) at Mar Chiquita Lagoon, Argentina. Ornitología Neotropical22:453-457
García Go, becKer Ph y Favero M (2011) Kleptoparasi- tism during courtship in Sterna hirundo and its re- lationship with female reproductive performance. Journal of Ornithology 152:103-110
García Go, becKer Ph y Favero M (2013) Intraspecifickleptoparasitism improves chick growth and re- productive output in Common Terns (Sterna hirun- do). Ibis 155:338-347
García Go, castano Mv, Paz ja, Paterlini ca, zuMPano F,y Favero M (2021) Hematologic metrics from olrog’s gull (Larus atlanticus) during the nonbreeding sea- son in Argentina. Journal of Zoo and Wildlife Medicine52:348-356
Garcia Go, Favero M y Mariano-jelicich r (2008) Red‐ gartered Coot Fulica armillata feeding on the grap- sid crab Cyrtograpsus angulatus: advantages and disadvantages of an unusual food resource. Ibis150:110-114
García Go, Favero M y vassallo ai (2010) Factors affec- ting kleptoparasitism by gulls in a multi-species seabird colony. The Condor 112:521-529
García Go, Favero M. y vassallo ai (2012) Interspecifickleptoparasitism by Brown-headed Gulls (Chroico- cephalus maculipennis) on two hosts with different foraging strategies: a comparative approach. Emu112:227-233
García Go, nicolli ar, castano Mv, zuMPano F, Favero M y biondi lM (2019a) Evaluation of neophobia in a threatened seabird: Olrog’s Gull (Larus atlanticus) as a case study. Emu 119:166-175
García Go, Paterlini ca, castano Mv, nicolli ar, zuMPa-no F y Favero M (2019b) Serum biochemistry values for the olrog’s gull (Larus atlanticus): a study based on free-living individuals. Journal of Zoo and Wildlife Medicine 50:330-336
García Go, Paterlini ca, Favero M, becKer Ph y bouwhuiss (2020) Age-, sex-and tactic specific kleptoparasi- tic performance in a long-lived seabird. Journal of Ornithology 161:183-188
Gatto a, Quintana F y yorio P (2008) Feeding behavior and habitat use in a waterbird assemblage at a ma- rine wetland in coastal Patagonia, Argentina. Water- birds 31(3): 463-471.
Giraldeau la y caraco t (2000) Social Foraging Theory.Princeton University Press, Princeton.
GóMez-tejedor h y de loPe F (1995) Cleptoparasitismo en la Gaviota Sombría (Larus fuscus): selección de víctimas y diferencias entre clases de edad. Ardeola42: 77-81.
Goss‐custard jd, cayFord jt y lea sG (1999) Vigilance during food handling by Oystercatchers Haemato- pus ostralegus reduces the chances of losing prey to kleptoparasites. Ibis 141:368-376
herrera G, Punta G y yorio P (2005) Diet specializationof Olrog’s Gull Larus atlanticus during the breeding season at Golfo San Jorge, Argentina. Bird Conserva- tion International 15:89-97
iribarne o, bortolus a y botto F (1997) Between-habitatdifferences in burrow characteristics and trophic modes in the southwestern Atlantic burrowing crab Chasmagnathus granulata. Marine Ecology Progress Se- ries 155:137-145
isla F (1997) Seasonal behaviour of Mar Chiquita ti-dal inlet in relation to adjacent beaches, Argentina.Journal of Coastal Research 13:1221-1232
josens Ml, Favero M y bo Ms (2010) Foraging ecology of the Great grebe Podicephorus major in Mar Chiquita lagoon (Buenos Aires, Argentina). Ardeola 57: 133-141
Krause j y ruxton Gd (2002) Living in groups. OxfordUniversity Press, Oxford.
Martin P y bateson P (1993) Measuring Behaviour, an In- troductory Guide. Second Edition. Cambridge Univer- sity Press, Cambridge
Mayds y aves arGentinas (Ministerio de Ambiente y
Desarrollo Sustentable y Aves Argentina) (2017) Categorización de las Aves de la Argentina (2015) Informe del Ministerio de Ambiente y Desarrollo Sustentable de la Nación y de Aves Argentinas, Bue- nos Aires, Argentina. [Edición electronica]
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Miller r, schiestl M, whiten a, schwab c y buGnyar t(2014) Tolerance and social facilitation in the fora- ging behaviour of free-ranging crows (Corvus corone corone; C. c. cornix). Ethology 120:1248-1255
Paredes r y insley sj (2010) Sex-biased aggression andmale-only care at sea in Brünnich’s Guillemots Uria lomvia and Razorbills Alca torda. Ibis 152(1): 48-62.
PuehrinGer-sturMayr v, stieFel t, Kotrschal K, Kleindor- Fer s y FriGerio d (2021) Social interactions change with season and age in Northern Bald Ibis. Journal of Ornithology 162(1): 277-288.
Quadri-adroGué a, seco Pon jP, García Go, castanoMv, coPello s, Favero M y MiGlioranza Ksb (2021) Chlorpyrifos and persistent organic pollutants in feathers of the near threatened Olrog’s Gull in sou- theastern Buenos Aires Province, Argentina. Envi- ronmental Pollution 272:115918
r core teaM (2021) R: A language and environment forstatistical computing. R Foundation for Statistical Computing, Vienna (URL https://www.R-project. org/)
ratcliFFe n, richardson d, scott rl, bond Pj, westlaKe c y stennett s (1997) Host selection, attack rates and success rates for Black-headed Gull kleptoparasi- tism of terns. Colonial Waterbirds 20:227-234
ravasi Mt, seco Pon jP, Paz ja, Favero M, y coPello s(2019). Use of winter habitat at an early age: spatial ecology and association with human activities of ju- venile Olrog’s Gull Larus atlanticus. Bird Conservation International 29:575-585
rothschild M y clay t (1952) Fleas, Flukes and Cuckoos:A Study of Bird Ectoparasites. Collins Son and Co. Ltd., Londres
shealer da, sPendelow ja, hatField js y nisbet ic (2005)The adaptive significance of stealing in a marine bird and its relationship to parental quality. Beha- vioral ecology 16:371-376
sieGel aF (1979) The noncentral chi-squared distri- bution with zero degrees of freedom and testing for uniformity. Biometrika 66(2): 381-386
sPivaK e, anGer K, luPPi t, bas c y isMael d (1994) Dis-
tribution and habitat preferences of two grapsid crab species in Mar Chiquita Lagoon (Province of Buenos Aires, Argentina). Helgoländer Meeresunter- suchungen 48:59-78
steele wK y hocKey Pa (1995) Factors influencing
rate and success of intraspecific kleptoparasi- tism among kelp gulls (Larus dominicanus). The Auk112:847-859
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Factors affecting the individual foraging behavior in a threatened seabird: Olrog’s Gull (Larus atlan- ticus) as a case study. Canadian Journal of Zoology99:658-664

How to cite

Francisco Zumpano, Germán Oscar García (2022). Intra-specific interactions during group feeding in the Olrog’s Gull (Larus atlanticus). El Hornero, Vol. 37 No. 1 (2022), 51-63. https://doi.org/10.56178/eh.v37i1.361

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[1] aGresti a (2002) Categorical data analysis. Second edi- tion. Wiley, Nueva York.

[2] aMat ja (1990) Food usurpation by waterfowl and wa-ders. Wildfowl 41:107-116

[3] annorbah nnd y holbech lh (2012) Relative abundan- ce, agonistic behaviour, and resource partitioning among three scavenging bird species in Ghana. Ma- limbus 34:1-8

[4] azPiroz ab, alFaro M y jiMénez s (2012) Lista Roja delas Aves del Uruguay. Una evaluación del estado de conservación de la avifauna nacional con base en los criterios de la Unión Internacional para la Con- servación de la Naturaleza. Dirección Nacional de Medio Ambiente, Montevideo

[5] barnard cj (1984) The evolution of food-scroungingstrategies within and between species. Pp. 95-126 en: barnard cj (ed) Producers and Scroungers: Stra- tegies of Exploitation and Parasitism. Croom Helm, Londres

[6] bates d, Mächler M, bolKer b y walKer s (2015) FittingLinear Mixed-Effects Models Using lme4. Journal ofStatistical Software 67(1): 1-48

[7] bautista lM, alonso jc y alonso ja (1998) Foraging site displacement in common crane flocks. Animal Be- haviour 56:1237-1243

[8] beauchaMP G (2017) Disentangling the various mecha-nisms that account for the decline in vigilance with group size. Behavioural Processes 136:59-63

[9] beauchaMP G (2019) On how risk and group size in-teract to influence vigilance. Biological Reviews94:1918-1934

[10] berón MP, García Go, luPPi t y Favero M (2011) Age-re- lated prey selectivity and foraging efficiency of Olrog’s Gulls (Larus atlanticus) feeding on crabs in their non-breeding grounds. Emu 111:172-178

[11] bertellotti nM y yorio P (2000) Age-related feedingbehaviour and foraging efficiency in Kelp Gulls La- rus dominicanus attending coastal trawlers in Argen- tina. Ardea 88(2): 207-214

[12] birdliFe international (2021) Species factsheet: Larus atlanticus (URL http://www.birdlife.org)

[13] boschi ee (1964) Los crustáceos decápodos Brachyu- ra del litoral bonaerense (R. Argentina). Boletín del Instituto de Biología Marina 6:1-99

[14] bower KM (2003). When to use Fisher’s exact test.

[15] American Society for Quality, Six Sigma Forum Magazi-ne 2(4): 35-37.

[16] brocKMann hj y barnard cj (1979) Kleptoparasitism in birds. Animal behaviour 27:487-514

[17] burGer j (1988) Foraging behavior in gulls: differen-ces in method, prey, and habitat. Colonial Waterbirds9-23.

[18] burGer j y GochFeld M (1979) Age differences in Ring-billed Gull kleptoparasitism on Starlings. The Auk 96(4): 806-808.

[19] castano M, biondi lM, zuMPano F, Favero M y García Go

[20] (2020) Behavioral responses to a novel feeding pro- blem in the Olrog’s Gull Larus atlanticus. Marine Or- nithology 48:149-155

[21] coPello s y Favero M (2001) Foraging ecology of Olrog’sGull Larus atlanticus in Mar Chiquita Lagoon (Bue- nos Aires, Argentina): are there age-related diffe- rences? Bird Conservation International 11:175-188

[22] crawley Mj (2007) The R Book. John Wiley and Sons,UK.

[23] duriez o, herMan s y sarrazin F (2012) Intra-specific competition in foraging Griffon Vultures Gyps ful- vus: 2. The influence of supplementary feeding ma- nagement. Bird Study 59:193-206

[24] escalante r (1966) Notes on the Uruguayan popula-tion of Larus belcheri. The Condor 68:507-510

[25] Fasano jl, hernández Ma, isla Fi y schnacK ej (1982) Aspectos evolutivos y ambientales de la laguna Mar Chiquita (provincia de Buenos Aires, Argentina). Oceanológica Acta 4:285-292

[26] Favero M (1996) Foraging ecology of Pale-Faced Shea-thbills in colonies of southern Elephant Seals at King George Island, Antarctica. Journal of Field Or- nithology 67:292-299

[27] Favero M, bachMann s, coPello s, Mariano-jelicich r, sil-va MP, Ghys M, KhatchiKian ce y Mauco l (2001) Aves marinas de sudeste Bonaerense. Pp. 251-267 en: iribarne o (ed) Reserva de Biosfera Mar Chiquita: características físicas, biológicas y ecológicas. Edi- torial Martín, Mar del Plata

[28] Flower tP y Gribble M (2012) Kleptoparasitism by at-tacks versus false alarm calls in fork-tailed dron- gos. Animal Behaviour 83:403-410

[29] Flower tP, child MF y ridley ar (2013) The ecologicaleconomics of kleptoparasitism: pay‐offs from self‐ foraging versus kleptoparasitism. Journal of Animal Ecology 82:245-255

[30] Furness rw (1987) Kleptoparasitism in seabirds. Pp.

[31] 77-99 en: croxall jP (ed) Seabirds: Feeding Biology and Role in Marine Ecosystems. Cambridge Univer- sity Press, Cambridge

[32] GaGlio d, sherley rb, cooK tr, ryan PG y Flower t(2018) The costs of kleptoparasitism: a study of mixed-species seabird breeding colonies. Behavio- ral Ecology 29:939-947

[33] García Go y biondi lM (2011) Kleptoparasitism by theCaracara Chimango (Milvago chimango) on the Ame- rican Oystercatcher (Haematopus palliatus) at Mar Chiquita Lagoon, Argentina. Ornitología Neotropical22:453-457

[34] García Go, becKer Ph y Favero M (2011) Kleptoparasi- tism during courtship in Sterna hirundo and its re- lationship with female reproductive performance. Journal of Ornithology 152:103-110

[35] García Go, becKer Ph y Favero M (2013) Intraspecifickleptoparasitism improves chick growth and re- productive output in Common Terns (Sterna hirun- do). Ibis 155:338-347

[36] García Go, castano Mv, Paz ja, Paterlini ca, zuMPano F,y Favero M (2021) Hematologic metrics from olrog’s gull (Larus atlanticus) during the nonbreeding sea- son in Argentina. Journal of Zoo and Wildlife Medicine52:348-356

[37] Garcia Go, Favero M y Mariano-jelicich r (2008) Red‐ gartered Coot Fulica armillata feeding on the grap- sid crab Cyrtograpsus angulatus: advantages and disadvantages of an unusual food resource. Ibis150:110-114

[38] García Go, Favero M y vassallo ai (2010) Factors affec- ting kleptoparasitism by gulls in a multi-species seabird colony. The Condor 112:521-529

[39] García Go, Favero M. y vassallo ai (2012) Interspecifickleptoparasitism by Brown-headed Gulls (Chroico- cephalus maculipennis) on two hosts with different foraging strategies: a comparative approach. Emu112:227-233

[40] García Go, nicolli ar, castano Mv, zuMPano F, Favero M y biondi lM (2019a) Evaluation of neophobia in a threatened seabird: Olrog’s Gull (Larus atlanticus) as a case study. Emu 119:166-175

[41] García Go, Paterlini ca, castano Mv, nicolli ar, zuMPa-no F y Favero M (2019b) Serum biochemistry values for the olrog’s gull (Larus atlanticus): a study based on free-living individuals. Journal of Zoo and Wildlife Medicine 50:330-336

[42] García Go, Paterlini ca, Favero M, becKer Ph y bouwhuiss (2020) Age-, sex-and tactic specific kleptoparasi- tic performance in a long-lived seabird. Journal of Ornithology 161:183-188

[43] Gatto a, Quintana F y yorio P (2008) Feeding behavior and habitat use in a waterbird assemblage at a ma- rine wetland in coastal Patagonia, Argentina. Water- birds 31(3): 463-471.

[44] Giraldeau la y caraco t (2000) Social Foraging Theory.Princeton University Press, Princeton.

[45] GóMez-tejedor h y de loPe F (1995) Cleptoparasitismo en la Gaviota Sombría (Larus fuscus): selección de víctimas y diferencias entre clases de edad. Ardeola42: 77-81.

[46] Goss‐custard jd, cayFord jt y lea sG (1999) Vigilance during food handling by Oystercatchers Haemato- pus ostralegus reduces the chances of losing prey to kleptoparasites. Ibis 141:368-376

[47] herrera G, Punta G y yorio P (2005) Diet specializationof Olrog’s Gull Larus atlanticus during the breeding season at Golfo San Jorge, Argentina. Bird Conserva- tion International 15:89-97

[48] iribarne o, bortolus a y botto F (1997) Between-habitatdifferences in burrow characteristics and trophic modes in the southwestern Atlantic burrowing crab Chasmagnathus granulata. Marine Ecology Progress Se- ries 155:137-145

[49] isla F (1997) Seasonal behaviour of Mar Chiquita ti-dal inlet in relation to adjacent beaches, Argentina.Journal of Coastal Research 13:1221-1232

[50] josens Ml, Favero M y bo Ms (2010) Foraging ecology of the Great grebe Podicephorus major in Mar Chiquita lagoon (Buenos Aires, Argentina). Ardeola 57: 133-141

[51] Krause j y ruxton Gd (2002) Living in groups. OxfordUniversity Press, Oxford.

[52] Martin P y bateson P (1993) Measuring Behaviour, an In- troductory Guide. Second Edition. Cambridge Univer- sity Press, Cambridge

[53] Mayds y aves arGentinas (Ministerio de Ambiente y

[54] Desarrollo Sustentable y Aves Argentina) (2017) Categorización de las Aves de la Argentina (2015) Informe del Ministerio de Ambiente y Desarrollo Sustentable de la Nación y de Aves Argentinas, Bue- nos Aires, Argentina. [Edición electronica]

[55] Medina al, tárano z y levin le (2008) Caracterizaciónde las posturas de amenaza en aves que compiten por el alimento. Memorias del Instituto de Biología Ex- perimental 5: 173-176

[56] Miller r, schiestl M, whiten a, schwab c y buGnyar t(2014) Tolerance and social facilitation in the fora- ging behaviour of free-ranging crows (Corvus corone corone; C. c. cornix). Ethology 120:1248-1255

[57] Paredes r y insley sj (2010) Sex-biased aggression andmale-only care at sea in Brünnich’s Guillemots Uria lomvia and Razorbills Alca torda. Ibis 152(1): 48-62.

[58] PuehrinGer-sturMayr v, stieFel t, Kotrschal K, Kleindor- Fer s y FriGerio d (2021) Social interactions change with season and age in Northern Bald Ibis. Journal of Ornithology 162(1): 277-288.

[59] Quadri-adroGué a, seco Pon jP, García Go, castanoMv, coPello s, Favero M y MiGlioranza Ksb (2021) Chlorpyrifos and persistent organic pollutants in feathers of the near threatened Olrog’s Gull in sou- theastern Buenos Aires Province, Argentina. Envi- ronmental Pollution 272:115918

[60] r core teaM (2021) R: A language and environment forstatistical computing. R Foundation for Statistical Computing, Vienna (URL https://www.R-project. org/)

[61] ratcliFFe n, richardson d, scott rl, bond Pj, westlaKe c y stennett s (1997) Host selection, attack rates and success rates for Black-headed Gull kleptoparasi- tism of terns. Colonial Waterbirds 20:227-234

[62] ravasi Mt, seco Pon jP, Paz ja, Favero M, y coPello s(2019). Use of winter habitat at an early age: spatial ecology and association with human activities of ju- venile Olrog’s Gull Larus atlanticus. Bird Conservation International 29:575-585

[63] rothschild M y clay t (1952) Fleas, Flukes and Cuckoos:A Study of Bird Ectoparasites. Collins Son and Co. Ltd., Londres

[64] shealer da, sPendelow ja, hatField js y nisbet ic (2005)The adaptive significance of stealing in a marine bird and its relationship to parental quality. Beha- vioral ecology 16:371-376

[65] sieGel aF (1979) The noncentral chi-squared distri- bution with zero degrees of freedom and testing for uniformity. Biometrika 66(2): 381-386

[66] sPivaK e, anGer K, luPPi t, bas c y isMael d (1994) Dis-

[67] tribution and habitat preferences of two grapsid crab species in Mar Chiquita Lagoon (Province of Buenos Aires, Argentina). Helgoländer Meeresunter- suchungen 48:59-78

[68] steele wK y hocKey Pa (1995) Factors influencing

[69] rate and success of intraspecific kleptoparasi- tism among kelp gulls (Larus dominicanus). The Auk112:847-859

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