Spatial Ecology of Migratory Passerines on Temperate and Boreal Forest Breeding Grounds

THE USE OF space reflects individual access to resources that are critical to survival and reproduction. Consequently, space use should be sensitive to a range of factors that have the potential to profoundly influence populations (e.g., Whitaker et al. 2007). The territory is one measure that has been employed to assess avian space use in the context of access to resources. In her seminal paper entitled “The Role of Territory in Bird Life,” Margaret Morse Nice (1941) synthesized a then long-running discussion on patterns of space use by birds into a functional classification system for the types of territories used by various species. Among these was her “Type A” territory, which referred to a defended area used for mating, nesting, and as a feeding site for young; such territories are maintained by most breeding passerines and other non-colonial bird species that have socially monogamous breeding systems (Nice 1941). The territory has since been widely accepted as the fundamental unit of space use for most passerine species (e.g., Morse 1989, Barg et al. 2005, Fisher and Davis 2010), and the concept of the home range, which may include both defended and undefended areas, has been notably absent from many discussions of passerine spatial ecology (e.g., Cody 1981, Haila et al. 1996, Cornell and Donovan 2010a; see also many species accounts in Poole 2005). In large part, this is because technological limitations required, until recently, that most studies of space use employ passive methods, such as territory mapping, that focus on the visual observation of prominently displaying males (Bibby et al. 2000). However, this focus may have led to an incomplete understanding of space use if individuals make extensive movements beyond surveyed territorial areas into a broader home range, are nonterritorial (e.g., during the postbreeding period), or behave cryptically or are not equally detectable throughout the area they frequent. Findings linked to recent technological progress and our growing knowledge of diverse aspects of avian ecology highlight the need to develop a more comprehensive understanding of passerine spatial ecology. Leading the way was the application of molecular genetics techniques to the study of avian breeding systems. This research revealed that extrapair mating was a regular occurrence in the vast majority of passerine species (Griffith et al. 2002), overturning assumptions that most socially monogamous birds were also sexually monogamous (Lack 1968). Because extrapair mates typically originate beyond territorial boundaries, this finding stimulated research into extraterritorial movements related to extrapair mating effort (e.g., Yezerinac and Weatherhead 1997, Stutchbury 1998, Woolfenden et al. 2005). During the same period, research in landscape ecology has revealed that individuals of many species are sensitive to habitat factors that occur beyond their territorial boundaries or at a scale larger than that of the territory (e.g., Hinsley et al. 1995; Betts et al. 2006, 2007). Similarly, ongoing research on other aspects of avian ecology, including dispersal, metapopulation dynamics, heterospecific attraction, and social information theory, has also led to greater understanding and recognition of the importance of extraterritorial movements by individuals, as well as interactions among birds distributed across a landscape (e.g., Hanski 1999, Walters 2000, Betts et al. 2008a). Field studies associated with these advances have been greatly facilitated by the development of miniaturized radiotransmitters, which allowed researchers to overcome many logistical hurdles that had hindered the collection of unbiased movement data. Use of radiotracking immediately demonstrated that individuals of some species make regular landscape-scale or extraterritorial movements (e.g., Hanski and Haila 1988, Williams 1990, Smiseth and Amundsen 1995), and studies that explicitly distinguish territorial areas defended by singing males from their more expansive home ranges are beginning to appear (e.g., Leonard et al. 2008, Anich et al. 2009). Information now available offers the hope of a more comprehensive and accurate understanding of the spatial ecology of breeding passerines. On the basis of recent research, we propose a model of space use in which individuals occupy defended territories embedded within potentially broader, multipurpose home ranges (Powell 2000) that are extensively traveled and for which they have a detailed cognitive map. However, there remains a substantial lack of basic knowledge regarding the overall spatial ecology of breeding passerines. Thus, our review focuses on recent research that provides insight into space use by passerines from the prebreeding through premigratory periods. Building on this, we offer general observations on the nature of passerine space use, assess the extent to which socially monogamous passerines use areas beyond the boundaries of their territories, and suggest some important future directions for research involving passerines and for conservation in temperate and boreal forests. Many migrants arrive at sites of higher latitude or altitude several weeks in advance of breeding (Benson and Winker 2001, Hahn et al. 2004, Hahn and Silverman 2007). Early arrival is associated with securing limited nest sites, high-quality territories, or mates (Francis and Cooke 1986, Cooper et al. 2009a), of and may be at this during and 2004, the was and at our study site in the of among the to arrive at our study were throughout the landscape and singing on territory during However, including was a regular occurrence in the area until and during such these to to on the at these of to individuals were the that birds were to on their breeding territories, was by that et al. 2009). has been to the ecology of birds during the prebreeding period, as this there may be on birds and may be important at this Hahn et al. see also Williams Consequently, patterns of space use during the prebreeding may in many important from in the of the breeding one that if individual birds the of such and use as then these areas and of their individual home ranges (Powell such sites may not be during with In birds available sites and a breeding territory, a and knowledge the breeding and of for use during future extrapair mating and Silverman 2006, Betts et al. an extensive cognitive of the landscape (e.g., and which this a potentially important for and information the of resources in the has been that birds for future breeding sites during the from the nest until on et al. some species for breeding sites until their breeding and The focus during such may well be the of both for the social that their regarding habitat et al. and for their potential future importance in species with breeding systems 1998, et al. During dispersal, an may well to the site and then from that to singing males as a et al. 2005, et al. 2005). the same social may be used by breeding during the and postbreeding et al. et al. 2005, Hahn and Silverman a of the various factors territories are not from on arrival until such as the arrival of in the to breeding et al. 2005). This prebreeding may several weeks et al. be on the of in arrival males and et al. that the habitat of during the prebreeding and breeding and et al. that the to territory in to of habitat may well be a during the prebreeding Betts et al. have in the prebreeding period, males from to sites as more information on habitat and other individuals sites as they also may also be that information is not immediately or available and to information by resources et al. or the of which are typically the more individuals that et al. that prebreeding defended territories within home ranges that were in area than in the breeding that habitat by as as were individuals that had not been in the study the prebreeding home ranges of to be This to as a space use during the prebreeding of space use during the is that the movements of breeding birds on their This a of to a that at a from the of nest through and and until the and the the researchers have employed as a to the spatial ecology of passerines during the (e.g., and studies have demonstrated to of spatial ecology, extraterritorial movements are a regular during the for both of many passerine species. individuals have been from to territory from their territories, from to of their territory, and in some extraterritorial and see also However, these extraterritorial have been referred to as (e.g., Stutchbury et al. which may and a that such movements are or than of defended territories and home and extraterritorial movement and and of territory for temperate and boreal passerines radiotracking during the prebreeding and breeding periods. not are in the of defended territories and home and extraterritorial movement and and of territory for temperate and boreal passerines radiotracking during the prebreeding and breeding periods. not are in the The extent of movement has been by studies that the of extrapair such studies not include of movements by individuals, they that some birds of both their territory during the to extrapair of extrapair occupy a territory, many are or territories and some typically from et al. 1998, et al. 2006, et al. 2007). However, there are also of species in which the of extrapair by is and some may well be from or more et al. 2005, et al. 2009). with studies have limited areas and are extrapair at a scale greater than that of the most et al. 1996, Woolfenden et al. 2005). Consequently, they and larger movements may be as by et al. a that was by an extrapair that a territory extraterritorial movements such as these are an important of space use that the of studies of the and movements of breeding birds on passive techniques such as territory mapping, and to measure the and movements of breeding birds (e.g., et al. et al. 2000). on are or In such research, were to be as of if individuals were not by they may have been as the territory studies have that birds are typically and during extraterritorial movements (e.g., Williams 1990, et al. 1997, and which their and to of the of territorial and extraterritorial movement in studies that employ passive studies are to these and movements within a and study the of of birds and of the scale of movement The of to such extraterritorial movements with the scale at which such movements occur et al. et al. some of individuals have that individuals make extraterritorial regular on territories to from their et al. at our study area in of of of were of individuals that were from their territories Whitaker for extraterritorial movements in the breeding among and with of have to the of such movements in extrapair mating and of the recent on passerine spatial ecology has been at this (e.g., Stutchbury et al. 2005). However, extraterritorial movements also include is that most passerines in some individuals have been from sites to and Williams 1990, and Stutchbury potential of territory is the of socially information on the of habitat in the area et al. 1999, and 2001, Betts et al. 2008a). However, many extraterritorial movements are on for potential mates and extrapair mating 1998, and and our of the importance of this may be by the research this has the of may the potential of they are territory during their is both breeding for the and the potential to extrapair are and Stutchbury et al. 2005). of extrapair mating are et al. et al. 2010), various studies that influence this through their of which males they with et al. et al. and the nature of their movements and Amundsen and 2000, et al. et al. in some have been to extrapair mates that are of or higher than their social which that is an important 1988, et al. Stutchbury of some species may or males and social mates to assess their et al. 1997, et al. 1998, Stutchbury In males make extraterritorial et al. 2005, et al. both males and of many other species territories on and Amundsen et al. 1997, et al. However, extraterritorial movements of and to the period, males make such throughout the breeding and Stutchbury and Stutchbury 2001, et al. Similarly, and that were during the and of their the territories of other males during these and led to more extraterritorial which that the potential for these was to extrapair also et al. also that information on the of potential extrapair mates and assess the of males et al. In some this information they as in which make to extrapair territories at this and Amundsen However, also many extraterritorial movements during the period, their social mates on most which that may also individuals to their territories see also and 2004, et al. of defended territories and home and extraterritorial movement and and of territory for temperate and boreal passerines radiotracking during the prebreeding and breeding periods. not are in the of defended territories and home and extraterritorial movement and and of territory for temperate and boreal passerines radiotracking during the prebreeding and breeding periods. not are in the The that males of several species focus their extraterritorial on the territories of (e.g., et al. 1998, Stutchbury is of the and of information on individual cognitive were most to on other territories during nest by not their was nest in the and observations have been for and in which males make extraterritorial their is their is not 1988, et al. home range has been as a area an has a of during a a territory is an area within an home range which has or use (Powell 2000, 2001, et al. The observations make that the most model of space use for many temperate and boreal passerines is that occupy a defended territory, typically from to embedded within a larger multipurpose home range of to and This model is individuals not make extraterritorial in such the home range and territorial boundaries are one and the studies have both territory and that home ranges that on the of territories et al. 2008, Anich et al. 2009). in many of the studies the of the home range that the defended territory typically the home ranges of and of their This spatial for social and of social also individuals their home range to the and associated with territories and make more or more than the territories and home ranges is the range in the extent of space use that has been across individuals of various species. in and the territories were and larger than the territories, et al. 2008, Anich et al. see also and Stutchbury Similarly, the home ranges were larger than the home ranges in these species et al. 2008, Anich et al. 2009). This is also in the in space use that a individual may through the of a breeding that home ranges of on during to an of during the postbreeding Consequently, study of in space use to the of factors that populations also insight into diverse aspects of a ecology (e.g., and and Whitaker et al. 2007). studies have that are and 1997, Cooper et al. which the of in many breeding nonterritorial are to because of their nature and Stutchbury and their has been by a of et al. 2009). However, the of a may also be as of a of which with site or widely throughout a home range a defended territory et al. occupy one of that males may and a territory, these individuals may a of territory or extensively through a large home range, a and Stutchbury may be in of from one to the this may also be a for individuals that extrapair and potential mates by widely through a home range than a breeding territory and Cooper et al. that this may territories of in across the broader landscape at greater spatial than individuals that are to a breeding However, in some are more high-quality habitat and may within a area and 2007). to the period, limited information is available on the ecology and movements of passerines during the to et al. that are to an is to that the range of movement patterns be more than during the have that some of their breeding territories and are in other or of their breeding range and and recently, the application of to the study of postbreeding movements has demonstrated that individuals within a breeding a range of and more extensive movement patterns In some a of with on the breeding territory other from the area et al. 1998, et al. 2000, and In many typically (e.g., and Stutchbury 1997, and and in was that these the was the and Stutchbury of movement more diverse if one that occur and et al. through the postbreeding and that on their breeding territories, to habitat to their territories, to sites and the study area were for in which of individuals on their breeding home ranges during the postbreeding and to other areas et al. However, several of the individuals that to their breeding territories, which that these areas a of their home ranges at that This is to observations of in which individuals home ranges with of during the postbreeding 2005). passerines several during the postbreeding period, potentially including the of of an territory or home range, for future breeding sites, or and of for The need to these the diverse range of patterns of space use among individual of a species at this associated with and premigratory may some individuals to to more sites, if is during individuals may to and et al. 1999, and 2007). studies have linked patterns of postbreeding movement to the of individual on their territories and may are more to extensively or during the postbreeding et al. 1999, and 2001, Betts et al. 2008a). This the that for birds the postbreeding be important both for the of an territory et al. also as a to for future breeding sites et al. Betts et al. that many sites were during the postbreeding and in many of these sites were during the breeding and for and passerines in temperate and boreal during the are from to or most not are in the and for and passerines in temperate and boreal during the are from to or most not are in the with limited has been at the spatial ecology of and birds to et al. they have been typically on their territory for some and then to other areas of to a they of this to a of et al. 1998, et al. and movements by birds they from have been to associated with of and habitat to and use sites and et al. for future breeding sites and of that be used to to the territory the and and et al. In to et al. that is passerines are or at a some movements may because individuals that then in areas et al. However, that the of individual and associated with a through the postbreeding period, immediately typically and in the they as many as individuals Whitaker that these were is to that was not and by factors such as social or of of birds the need to for the of and to and premigratory for as well as to and 1989, et al. et al. studies have and extensive movements on a scale of such movements may be associated with these their et al. 1998, et al. and they may also be a to for sites that be used in the future (e.g., breeding to distinguish and have et al. 1998, et al. and some studies have in survival et al. 2006, et al. linked to the to and et al. 1998, and Stutchbury 2008, and However, et al. that birds movements and which the or in with with an that to the that they were a for suggest the importance of for and they not other on (e.g., and et al. the same they the that individuals are by a and a cognitive of the The observation that during the postbreeding many to in habitat the also that birds use postbreeding social to information on potential future breeding sites et al. Cornell and Donovan is well that many passerines to breeding territories, to nest (e.g., may also be the that individuals to extraterritorial sites used the and (e.g., and review of patterns of and breeding is beyond the of this paper and has been (e.g., and et al. However, in the is important to that if individuals occupy large home then a territory that within that home range an from to a site beyond the movements not in the of the a in a area or of a home range is Because they typically at of (e.g., and passerines are and make landscape-scale movements in This to range large areas to such resources as and mates and to information on habitat and the and of resources and in the there are potential associated with including and the of studies have that notably their use of habitat to and 1997, et al. 2001, survival that to is or at during the breeding in general et al. 2008, Whitaker et al. and during extraterritorial movements in and Stutchbury 2001, Woolfenden et al. 2005). are also of of during (e.g., Stutchbury et al. which that is in of and also et al. 1998, and 2000). Thus, in many the of extensive movement during the breeding may be from a nest or which to of nest nest or (e.g., and and of large areas, our review that an individual bird may several patterns of movement the of a breeding individuals in a employ patterns of space use at one This the importance of scale patterns of space use, which are and by a range of and factors during the of a breeding However, of this of spatial within the broader of a home range, if at an that or that be as within a may in use of of an overall home and that with their to the same sites in Similarly, individuals may make movements to social information during the postbreeding However, if they then use this cognitive future breeding sites et al. 1999, Betts et al. to these areas as of the home range 2000). this broader of passerine space use during the breeding period, is that and space use an The that most breeding passerines make extraterritorial movements during the that a of space use is This is overcome by to a more model in which breeding individuals typically a territory embedded within a larger home range that they for a of and for which they a detailed cognitive map. to or more within territories include territorial and (e.g., Betts et al. birds typically more territory (e.g., Stutchbury However, of a territory may occur during of the breeding typically the prebreeding and and many individuals one or more throughout the of a breeding this a to site within the breeding which an individual (e.g., et al. 1999, 2005). In other individuals make including dispersal, and (e.g., and et al. 1998, et al. 2009). and these of space use a of the range of patterns of movement and of the scale in which they are may the and 1990, 2000, et al. highlight the importance of to home ranges and other used The has been widely employed (e.g., and 2001, and is sensitive to large areas of and offers information on the of within the home range and researchers should home ranges in of which a used area to be in of the of in and 1996, Barg et al. 2005). of techniques have been and have been (Powell 2000, 2001, et al. In the has has many and 1996, and is well for use with passerines (e.g., Barg et al. 2005, Leonard et al. 2008, Anich et al. 2009). However, these are individuals are not in which a is (e.g., et al. a of information on the spatial ecology of passerines has been in the that had been to in extrapair mating the that beyond territorial studies have on since that and this research substantial in of most there are studies to our knowledge of movements have also been studies of and a of extrapair mating et al. Woolfenden et al. 2005, et al. the to are et al. 2010), as are many of the the that the use during focus on of habitat use through of displaying males (e.g., Whitaker and may also that important habitat of have been in the development of conservation across a range of studies on temperate and boreal one of the most and aspects of space use is the in the scale of This is one individuals during a or a individual of the breeding the of passerines to the extent of their movements to their and and The range of this to for habitat or is critical to as this is one of the that birds employ to to et al. 1997, Walters the that such (e.g., et al. 1997, Whitaker et al. Leonard et al. and their associated (e.g., a to may a greater understanding of the of to by species. is a growing of that landscape-scale habitat the movement of passerines. and have been to make more extensive movements in than in temperate and boreal and 2001, and Stutchbury 2001, and Stutchbury 2004, Anich 2008, Leonard et al. However, to the of habitat species to habitat for a of in this (e.g., et al. 2001, and 2002), and a range of other aspects of an as well as and Consequently, individuals use these may be of habitat (e.g., Hinsley 2000, and 2002), and there may be of habitat or beyond which individuals (e.g., Betts et al. Cornell and Donovan may be to these to the movement and of individual extraterritorial may be more the habitat is or of such that males home ranges in may need to habitat to and Stutchbury 2004, Leonard et al. This a and and or of 2000, and Stutchbury et al. 2009). important to conservation is an understanding of landscape-scale habitat that from more of individual space During the period, home ranges of passerines may be to more than larger than territories et al. 2008, Anich et al. 2009). This greater for individuals, or if one the and postbreeding movements of Use of large home in some (e.g., and Stutchbury 2004, offers a to the landscape-scale habitat in many species of (e.g., Hinsley et al. and 2005, Betts et al. of territories is by habitat use the broader home range that an individual is also critical for understanding the habitat ecology. is important that we the of home ranges within and of territories within home to for habitat However, studies of passerines have explicitly habitat at the scale or habitat in territories with that in broader home the potential importance of extraterritorial habitat for at some species in of and extrapair mating may be by the of habitat at the the in which and use the broader landscape has to both individual survival and future et al. of the that on extrapair mating extraterritorial movements as or on the territories of However, individuals of most species make regular extraterritorial movements into a broader landscape for which they have detailed knowledge and a cognitive and which they may for a of throughout the breeding Consequently, more and accurate to this area as an multipurpose home range than a landscape to be This the from a of as in and territories from to one in which in have home defended territories, have a detailed knowledge of their and with also have a of the and of resources across the broader an area that for some species may several This as an for diverse related to the and of populations and of individuals across well to studies of in landscape-scale habitat (e.g., be as in the is also with extrapair mating systems and the which has been to of territories in some species of 1998, et al. to a more comprehensive understanding of metapopulation and et al. diverse types of information the to a cognitive and are used in future This was greatly as a of from and

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