Sibling Rivalry PSYCH 3T03 Lecture 17 Animal Behavior – Chapters 10 & 11 1. What is sibling rivalry? 2. Why is there conflict between siblings? 3. Facultative vs obligate strategies 4. Lethal & non-lethal competition 5. What behaviours have evolved as a consequence? Blue footed booby © Derek Keats Reproductive Conflicts & Trade-offs onf c l a u Sex Pa Each individual ‘wants’ to maximise its own lifetime reproductive fitness lict prin s f f -O r ent g Optimal strategies will differ between males – females – offspring Uneven parental investment creates conflict: One parent vs the other valr i r g n Sibli y One breeding season vs another One sibling vs another Benefits of Parental Care Increases the survival and fitness of offspring That could mean: Greater survival to adult Faster growth Healthier / better able to attract mates Social development & learning Indirect benefits to parents Paper wasp © Alvesgaspar Trade offs in Parental Care When should a parent stop parental care and invest in further reproduction? Greater investment = greater costs to parents But for offspring greater investment = greater benefits Potential conflict over: 1. Duration of parental care 2. Distribution of parental care © Alex Proimmos Examples of Parent-Offspring Conflict 1. Resource provision Offspring want more food 2. Defence Risky behaviours for parents benefit offspring 3. Clutch size More offspring good for parents but more siblings may be bad for offspring 4. Sex ratios Optimal sex ratio of offspring may be different to optimal sex ratio of siblings Parents want to maximise reproductive output… Examples of Sibling-Sibling Conflict … offspring want to maximise individual investment 1. Resource provision Offspring want more food from parents 2. Inheritance Offspring want access to natal territory 3. Future rivals Siblings are competition for future territory/food 4. Sex ratios Future competition with same sex siblings for mates Direct and Indirect Competition Offspring may compete for attention or actively interact with each other 1. Begging 2. Scrambling 3. Sublethal dominance 4. Siblicide © Katie McVey, USFWS Competition for resources in cichlid fish Territorial fish found in Lake Tanganyika Variabilichromis moorii Biparental care – defence but no provisioning 1. © JBenoit Offspring compete for access to food within natal territory Competition for resources in cichlid fish Territorial fish found in Lake Tanganyika Satoh et al. 2021. Behav Ecol. Biparental care – defence but no provisioning 1. Offspring compete for access to food within natal territory 2. Larger juveniles monopolise food and grow more quickly than subordinates Competition for resources in cichlid fish Territorial fish found in Lake Tanganyika Satoh et al. 2021. Behav Ecol. Biparental care – defence but no provisioning 1. Offspring compete for access to food within natal territory 2. Larger juveniles monopolise food and grow more quickly than subordinates 3. Subordinates leave nest earlier to find food Dominant juveniles get more access to food And receive parental care for longer Competition for future territory in jays Socially breeding bird found in northern Eurasia Siberian Jay - Perisoreus infaustus Biparental care with previous offspring helping 1. Offspring can either disperse after fledging or remain in the natal territory 2. Dominant offspring have better access to food within the territory © Bouke ten Cate Competition for future territory in jays Socially breeding bird found in northern Eurasia Ekman et al. 2002. Anim Behav. Biparental care with previous offspring helping 1. Offspring can either disperse after fledging or remain in the natal territory 2. Dominant offspring have better access to food within the territory 3. Dominant offspring (larger size) more likely to remain in natal territory Competition for future territory in jays Socially breeding bird found in northern Eurasia Ekman et al. 1999. PRSB. Biparental care with previous offspring helping 1. Offspring can either disperse after fledging or remain in the natal territory 2. Dominant offspring have better access to food within the territory 3. Dominant offspring (larger size) more likely to remain in natal territory 4. Remaining offspring have more breeding success over their lifetimes Parental insurance policies Parents may produce more offspring than they can look after Not all offspring will survive Back up option just in case Not all offspring are equally fit Competition allows for selection of strongest Conditions change Often only a short period of time in which to breed © Diego Cue Overproduce early and reduce numbers is a better strategy than waiting to catch up later Siblicide Offspring may kill their siblings Reduce competition for parental care Remove future rivals Directly acquire resources (cannibalism) Maybe direct (killing) or indirect (out competing) Maybe obligate or facultative Earwig © Photo by Tom Oates, 2010 Siblicide provides extra resources Earwigs raise offspring in communal nests Forficula auricularia Nymphs do not disperse until after they molt 1. © Tom Oates, 2010 Nymphs will kill and cannibalize siblings Siblicide provides extra resources Earwigs raise offspring in communal nests Dobler & Kölliker. 2009. Behav Ecol. Nymphs do not disperse until after they molt 1. Nymphs will kill and cannibalize siblings 2. Survivors benefit from Reduced competition Nutritional gain from eating sibling 3. Cannibalism more common among halfsiblings compared to full-siblings © Tom Oates, 2010 Siblicide provides extra resources Intrauterine cannibalistic siblicide in sand tiger sharks Carcharias taurus Sand tiger sharks give birth to live young 1. © Shankar S Female will produce many embryos that start development Siblicide provides extra resources Intrauterine cannibalistic siblicide in sand tiger sharks Chapan et al. 2013. Biol Lett. Sand tiger sharks give birth to live young 1. Female will produce many embryos that start development 2. Eggs develop and hatch within the mother 3. First hatchling to hatch eats all of its siblings and undeveloped eggs 4. Growth rate increases massively Siblicide skews the sex ratio Future reproductive success can depend on the sex ratio Copidosoma floridanum Parasitic wasp with haplodiploidy system Two types of offspring Metamorphosing reproducing offspring Non-metamorphosing offspring with large jaws © John Rosenfeld, Bugguide.net Siblicide skews the sex ratio Future reproductive success can depend on the sex ratio Grbić et al. 1992. Nature. Parasitic wasp with haplodiploidy system Two types of offspring Metamorphosing reproducing offspring Non-metamorphosing offspring with large jaws Clonal non-metamorphosing offspring kill males One male can fertilize all females Killing brothers gives more resources to sisters © John Rosenfeld, Bugguide.net Competition can lead specialised adaptations Competition between siblings can drive the evolution of morphological weapons Domestic pigs Sows typically produce 20% more offspring than they can raise Competition among piglets for access to milk Despite not needing teeth to drink milk Piglets born with fully erupted canines Teeth used to fight siblings Often removed by farmers to protect their livestock Competition can lead specialised adaptations Competition between siblings can drive the evolution of morphological weapons Fraser & Thompson. 1991. Behav Ecol Sociobiol. Experimentally removed the teeth of the heaviest piglets from multiple litters Compared growth rate between piglets with and without canine teeth Piglets with canines grew faster Effect greatest in large litters Where competition was most intense Resource availability Facultative siblicide in hyena Spotted hyena frequently give birth to twins Crocuta crocuta Twins born regardless of environmental conditions 1. Competition between siblings will depend on how much food the group can provide Facultative siblicide in hyena Spotted hyena frequently give birth to twins Hofer & East. 2008. Behav Ecol Sociobiol. Twins born regardless of environmental conditions 1. Competition between siblings will depend on how much food the group can provide If cubs are growing slowly rate of siblicide increases Facultative siblicide in hyena Spotted hyena frequently give birth to twins Hofer & East. 2008. Behav Ecol Sociobiol. Twins born regardless of environmental conditions 1. Competition between siblings will depend on how much food the group can provide If cubs are growing slowly rate of siblicide increases Total food provision does not decrease after siblicide Growth rate of surviving cub increases after siblicide Facultative vs Obligate Siblicide Blue Footed Booby Masked Booby © MC VC © pjd1 Facultative vs Obligate Siblicide Blue Footed Booby Both species produce two eggs and incubate immediately Asynchronous hatching = alpha and beta chicks differ in size Blue footed booby is facultative © MC VC Only kills sibling when food is scarce Masked Booby Masked booby is obligate Always kills sibling immediately © pjd1 When conditions are favourable Blue Footed can raise two chicks, but Masked cannot Control of Facultative vs Obligate Siblicide Great Blue Heron Great Egret © Russ © Mike Baird Control of Facultative vs Obligate Siblicide Great Blue Heron Both species have similar breeding behaviour Great Blue Heron siblicide is rare © Russ Great White Egret Great White Egret siblicide is common Up to 85% of 4 egg clutches will have siblicide Chicks attack each other Why are there differences in the rate of siblicide? © Mike Baird Control of Facultative vs Obligate Siblicide Great Blue Heron Cross fostering experiment (Mock 1984. Science) Herons raised by egrets and vice versa Egret chicks in heron nests could not monopolise food © Russ Great White Egret High competition and siblicide but all chicks small and weak Heron chicks in egret nests increased aggression Increased siblicide among heron chicks Largest chicks monopolized food Important role of parent behaviour © Mike Baird The role of parents – competition a benefit? Parents can encourage or discourage sibling competition 1. Selectively feed particular offspring 2. Preventing / ignoring aggression 3. Synchronous / asynchronous incubation – control age difference 4. Control the number of offspring produced / infanticide © Chanel City Camera Club Any Questions? PSYCH 3T03 Lecture 16 1. For offspring increased allocation to the detriment of parents and/or siblings can be an individual advantage 2. Siblings will compete for access to immediate benefits (e.g., food) and future opportunities (e.g., mating access) 3. Parents often produce more offspring than they can raise 4. Siblicide is common and can be beneficial for both parents and surviving offspring Great egret © Mike Baird