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Pair of female Northern Cardinals (Cardinalis cardinalis) rumbling CHAPTER 11: AVIAN SOCIAL BEHAVIOR Friday March 20 th going out for an extended lab time, no lecture SOCIAL BEHAVIOR This chapter focuses more on non-breeding social interactions Flocking species and acquisition of status in flocks Foraging or roosting spots Territoriality Defending resources from others A female Eastern Bluebird (Sialia sialis) attacking a female House Sparrow (Passer domesticus) over a nest box Interactions with other species entirely Food, territory, other resources SOCIAL BEHAVIOR Individual spacing and ‘personal space’ important SOCIAL BEHAVIOR Territoriality Defending a fixed area for your personal use Can be temporary or longer term Contains resources that you use exclusively Food, nest sites, roosting sites, safe areas from predators May defend from just your own species, may defend from others as well Neighbors and floaters – the threat differs in many ways Breeding territories, wintering territories, general purpose territories Defense or announcement of ownership by songs, calls, or displays SOCIAL BEHAVIOR Territory distribution of Northern Cardinals (Cardinalis cardinalis) at Aullwood Audubon Center, Dayton, OH Typical song posture of a male cardinal SOCIAL BEHAVIOR Benefits to territory defense Assuring a limited resource is available Costs to territory defense Must expend energy to keep your resource safe Chasing, fighting, displaying Golden-winged Sunbird (Nectarinia reichenowi) Payoff, protecting the energy source means energy to pay back expense SOCIAL BEHAVIOR Defending a territory from competitors, and defending the optimal size territory, is only feasible when the costs of doing so (energy, potential injury) are outweighed by the benefits (food, safety) When costs are too great territory is either abandoned (become non-territorial) or allowed to shrink (give up some until defense is feasible) SOCIAL BEHAVIOR Territory size required varies with species and with resource being used Larger species typically need more than smaller species (general space) Predators need more area than herbivores (energy dynamics) Quality of the resource on the territory can also determine size Archilochus colubris SOCIAL BEHAVIOR Acorn Woodpecker (Melanerpes formicivorus) granaries are a valuable resource, strongly defended by a group. SOCIAL BEHAVIOR Floaters and neighbors are always looking to take territory A territory owner has precedence over an intruder Owner knows the value and knows the lay of the land Experimentally remove an owner and usurpers fight to remain based on how long they’ve occupied territory Neighbors will take some of what’s yours if that is allowed Long-term territoriality, boundaries are established ‘Dear Enemy’ – ‘I know you and what you can do, you cannot steal from me’ ‘Nasty Neighbor’ – ‘I know you and what you can do and what you have, I can take it from you as you’re weak’ SOCIAL BEHAVIOR Within flocking species dominance status can be important Better access Better access Better access mates Better access territories to food to safety to future to future There are costs associated with this Some species, dominants have high corticosterone levels Red Junglefowl (Gallus gallus) females establish ‘pecking orders’ among their flocks for access to food and mates SOCIAL BEHAVIOR Social interactions can be what keeps order in groups and what keeps your resources yours, but Energy it takes to fight Costs due to injury – can be very expensive (death the most expensive) There must be other ways to communicate besides just jumping right to the attack! SOCIAL BEHAVIOR Physical appearance, ornamentation, can be used to quickly identify who is likely to be dominant and who isn’t. Ornament expression typically co-varies with age, quality, sex, or steroid hormones in such a way as to reliably indicate who is who. Harris's Sparrow, Zonotrichia querula “Stud” Not so much a ‘Stud’ SOCIAL BEHAVIOR Experimental modification of ornaments and status Harris’s Sparrows Only with implant of T could non-studs dyed to look like studs succeed House Sparrows Birds dyed to look dominant succeeded for a bit, then were tested Northern Cardinals A pale cardinal held his own even if he was super red originally Alteration of red didn’t change flock status, something other being used? SOCIAL BEHAVIOR Ocellated Antbird (Phaenostictus mcleannani) Bicolored Antbird (Gymnopithys leucaspis) Benefits of dominance within flocks, either of a single species or mixed species, that you get the best food. Not just ornaments but sheer physical size can be important Spotted Antbird (Hylophylax naevioides) SOCIAL BEHAVIOR Large Ocellated Antbirds (~50g) take precedent at the center of Army Ant swarms, here is the best insect foraging opportunities White-plumed Antbird (Pithys albifrons) SOCIAL BEHAVIOR Aggressive (agonistic) behavior A physical interaction is going to ensue, one will win and one will not If ornaments that co-vary with status not enough, fights may ensue Typically birds of similar status within flocks, or of similar ornamentation Often a stereotypical set of behaviors or postures to show intent Threat, displacement, attack/chase Bird that isn’t winning or isn’t escalating has appeasement display SOCIAL BEHAVIOR Don’t be fooled by group selection arguments, avoiding this is good for both the loser and the winner. This is what displays are trying to circumvent, here two male Yellow Warblers (Dendroica petechia) really battle it out SOCIAL BEHAVIOR Peeved and ready to do something about it Not ready to rumble, trying to diffuse the situation Steller's Jay (Cyanocitta stelleri) SOCIAL BEHAVIOR Ultimate attack stance in Northern Cardinals (Cardinalis cardinalis) SOCIAL BEHAVIOR Surprisingly some aspects of courtship have aggressive components Some suggest females incite males to fight to pick the best Males may need to overcome gut reaction to a conspecific A male could be mistakenly be trying to defend against an intruding female Mate has disappeared very recently SOCIAL BEHAVIOR Aggressive behavior makes one think of males, and males have been focused on Females are aggressive too Status in a flock/group Territory boundaries Access to males Defense of nesting resources SOCIAL BEHAVIOR Flocks – groups of birds, can be small (6 or so) can be huge (millions of individuals) Can be all the same species, can be comprised of multiple species – more later This has costs and benefits Benefits Safety in numbers, ease in finding food (work of off producers), more time devoted to foraging Costs Someone takes your food (scrounger), constant fighting for dominance, increased disease transmission, food might not be super -abundant SOCIAL BEHAVIOR Red-billed Quelea (Quelea quelea) flocks are enormous; promotes safety, and can manipulate the scrounger/producer system SOCIAL BEHAVIOR Young Common Ravens (Corvus corax) use flocks for information sharing and to gang up for resource acquisition SOCIAL BEHAVIOR Being in a flock can help you avoid predation (dilution effect, confusion effect, and more eyes on the look-out) Predator SOCIAL BEHAVIOR European (Common?) Starlings (Sturnus vulgarus) alter their behavior when they cannot easily see others, shows the benefits of associating in flock SOCIAL BEHAVIOR Multiple factors go into decisions concerning flock size, if dynamics around the flock change the optimal inclusion of more or less individuals will alter as well. Notice the cost of fighting with others over access to resources. SOCIAL BEHAVIOR Mixed species flocks – lots of species moving together Why do this? Some cases, the resource you’re after forces you together – antbird species following army ants Rare or territorial species may not have enough of their own around, joining with others a must (safety, better foraging) Surprisingly social signals are interpretable across species groups Appearance and behaviors associated with dominance cross species boundaries Often habitual associations lead to similar appearances SOCIAL BEHAVIOR Some cases individuals live or associate in super groups Colony nesters – enormous groups nesting near each other ~13% of species do this, primarily seabirds Shortage of safe nesting spots, everyone congregates where it’s best/safest Communal roosting Large groups offer safety – intimidating to predators and many eyes on the watch Large groups offer information – ‘Who ate well today, perhaps I’ll follow them tomorrow’ So many together, if sheltered, may reduce energy demands for thermoregulation May be easier to find a mate in the group But, energy spent on aggressive interactions over dominance and individual spacing; disease transmission SOCIAL BEHAVIOR Royal Tern (Thalasseus maximus) nesting colony, just area around body ‘secured’ SOCIAL BEHAVIOR Not really a communal roost here, but crowding can have detrimental impacts. Here, birds on the bottom of the piles have been known to suffocate and if one individual is sick, everyone is going to get it.