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Bird Territories: a Key to Understanding Bird Behavior Robert A Connecticut College Digital Commons @ Connecticut College Biology Faculty Publications Biology Department Spring 1987 Bird territories: A key to understanding bird behavior Robert A. Askins Connecticut College, [email protected] Follow this and additional works at: http://digitalcommons.conncoll.edu/biofacpub Part of the Ornithology Commons Recommended Citation Askins, R. A. 1987. Bird territories: A key to understanding bird behavior. American Birds 41: 35-40. This Article is brought to you for free and open access by the Biology Department at Digital Commons @ Connecticut College. It has been accepted for inclusion in Biology Faculty Publications by an authorized administrator of Digital Commons @ Connecticut College. For more information, please contact [email protected]. The views expressed in this paper are solely those of the author. BEHAVIOR Territories:a key to understandingbird behavior The role of amateur ornithologists in perplexingquestions arise with the ORarrivalTHE ofCAREFUL birds in FIELD spring.OBSERVER, Why do developing a major scientific theory. male Red-winged Blackbirds migrate two or three weeks before the brown, stripedfemales? Why do oriolessing in- cessantlyduring the first week of their arrival from the south? What could Robert A. Askins possibly motivate two flickers in a gar- den to flick their wings and bob their heads at each other, and then tumble together with flashing yellow wings? Scientific theories are most intriguing when they offer a single,simple expla- nation for such disparate phenomena, and the theory of territoriality has this advantage. The concept of territory providesinsights from which any ama- teur naturalist can benefit, since it ex- plainsmuch aboutthe daily behaviorof animals. Not surprisingly,the concept was originally developed by amateur naturalists, bird watchers whose pa- tience and skill for careful observation allowed them to detect a pattern in the freneticspring activity of birds. One of these bird watchers was Eliot Howard, an English businessmanwho lived in a house among pasturesand woodlands overlooking the River Sev- ern. Before leaving for work in Worces- ter. where he was director of a steel- manufacturing firm, Howard arosebe- fore daylight and spent the early morning hoursobserving the birds near his house. In 1920, after decades of watchingthe behaviorof birds,he wrote Territory in Bird LiJ& a book that greatly changedthe study of birds and other animals. Before this time, orni- thologistshad studied birds primarily by shooting them for their skins and collectingtheir eggsand nests,organiz- ing these into collectionsand naming Yellow-headedBlackbird males arrive in a breedingmarsh area befitreft,males. They evaluate the differentforms. This wasa necessary the habitat .forJood availabilio, and sql•' nest and roostsites. commit to a specificpatch an entireseason. and thereafterinvest heavily in defi,nse qf that territory.Because these decisions stagebecause it was essentialto impose havebeen made independento. f the presenceqffi, males. whenthey arrive; they are in a position some order and apply some labels to to assessthe quality or fitnessqf a male and his patch oJreal estateand to then choosea mate. the bewildering variety of thousandsof Males make habitat deciskmswhile fi•r ft,mal•:v, selecting a mate is equivalentto making a different kinds of birds. Howard, how- territory decision.Photo/S.R. Drennan. Volume 41, Number I 35 is such that males leave their winter feeding grounds for a few hours each day in late winter to claim an area and protect its boundaries. Defense of the territory becomesincreasingly impera- tive as the seasonadvances, and when a female joins the male, the pair coop- erate in defending their plot against other Reed Buntings. Song is used mainly to proclaim territory-ownership to other Reed Bunting males. Often males enter a fight in full song. In Howard's view, territoriality con- sistsof two components:(1) the occu- pancyof a particularplot of groundand (2) defenseof this areaagainst intrusion by membersof the same species(other than the mate). Territories vary greatly in size, from the squaremiles patrolled by a falcon to the half acre of willow thicket claimed by a warbler. Howard even consideredthe tiny areas around the nestsof cliff-nestingseabirds to be territories, since they are defended against other individuals of the same species. A pair of Common Murres Territorialencounter between Northern (Yellow-shafted) Flickers ( Colaptes auratus). Two female ( Uria aalge) or Razorbills(Alca torda) flickers (foreground)display aggressively near a nestcavity while a male watchespassively. defendsthe few squarefeet of cliff ledge During high-intensityterritorial displays, flickers face one anotherand bob their heads.At where their nest will be built. They de- the sametime they revealthe brightyellow on the undersideof the wingsand tail by raising fend only the nest site, not a food the wingsand spreadingand twistingthe tail (Lawrence1967}. Illustration/Don Luce. source, since they forage over broad areas of ocean. Similarly, male Ruffs ever, inspired many ornithologiststo marshes.Each male selecteda promi- (Philomachuspugnax) defend tiny areas study the biology of birds in a different nent position,the top of an alder or tall on a displayarena where males congre- way, in the field watchingthe activity reed, where it sangand preened. Early gate solelyto mate with females.How- of individual birds. His conceptionof in the seasonit often left thisspot, flying ard also considered these to be territo- the territory,an areadefended by a male a long distanceto the feedinggrounds. ties,and he probablywould haveagreed during the breeding season, demon- But as spring approached, these ab- with the simple, broad definition sug- stratedhow a usefuland generaltheory sencesbecame shorter and lessfrequent, gestedby Noble (1939): "territory is any could be derived from careful bird and soon the male Reed Bunting spent defended area." watching. all of its time at its selectedspot in the Howard's book awakened many or- Howard watched the Reed Buntings marsh. nithologiststo the phenomenon of ter- (Emberiza schoeniclus)that nestedin a While female Reed Buntings were ritoriality. How could such an appar- marshy meadow near his house. The still foraging together in winter flocks, ently obviousand predominant aspect Reed Bunting,an inhabitant of marshy the males spent more and more time of bird behavior have been overlooked edgesin Europe, is a small brown finch singingtheir tinkling song from their for so long? Actually many earlier au- with black head and throat. Howard special perches. If another male ap- thors had described territorial defense noticedthat in winter thesebuntings left proachedthe perch,he wasdriven away. in birds(Lack 1944).Aristotle described the wet meadows and reed-beds,where Fighting often occurred in such situa- the defenseof largeareas by ravensand they had spent the summer, to forage tions. The male was usually not joined eagles in Historia Animalium. The in open cultivated areas, often in the by a female until late March. The pair "walks" (apparentlyterritories) of semi- company of other speciesof finches. then drove other, neighboring pairs domesticMute Swans(Cygnus olor) are They normally returned to the marsh away from the area around the singing describedin 17th century Englishgame only at night, when groupsof buntings perch. Later, nestswere built and eggs laws,and many early ornithologistsde- roostedtogether low in the rushes.Cu- laid, and the frequencyof both singing scribed territorial behavior. In 1868, a rious about how these winter habits and fightingdeclined. German biologist, Bernard Altum, changedas springapproached, Howard Howard (1920) explained these wrote Der Vogel und sein Leben (The watched the birds through the cold events in terms of the need for a plot of Bird andHis Life), a bookthat describes morningsof February.In mid-February marsh, a territory, in which a pair of territoriality in birds in much the same he noticeda change:instead of joining Reed Buntingscould find a nestsite and detail and manner as Howard's Terri- flocks of finches in the usual feeding search out enough food to raise their tory in Bird Life (Mayr 1935).Howard's areas,male buntingsscattered across the young. The competition for territories discoveryof territoriality more than 50 36 AmericanBirds, Spring 1987 yearslater was apparentlyindependent, thoughout this period. The position of since he had not read Altum's book. his territory shiftedprogressively west- More important, it was Howard who ward by a total of 50 meters between convincedothers of the generalappli- 1930 and 1934, but later returned to its cability of territory theory and the use- original location. However, femalessel- fulnessof studyingthe behavior of in- dom remained with the same male from dividual birds. one year to the next; if they survived Following Howard's lead, many or- the winter, they normally appearedon nithologistswere inspiredto study ter- anotherterritory with a new male at the ritoriality in birdsduring the 1930sand beginningof the next breedingseason. 1940s. One of the most informative Occasionallythey even switchedmates studieswas done by another amateur, during the middle of the summer, and Margaret Nice, who spent eight years raisedtheir secondbrood of youngon studyingthe SongSparrows (Melospiza another territory. Both males and fe- melodia) around her house in Colum- males defended the territory, usually bus, Ohio. Nice was a well-educated
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