Breeding Dispersal and Philopatry in the Tree Swallow
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The Condor 106:768±776 q The Cooper Ornithological Society 2004 BREEDING DISPERSAL AND PHILOPATRY IN THE TREE SWALLOW DAVID W. W INKLER1,6,PETER H. WREGE1,PAUL E. ALLEN2,TRACEY L. KAST2, PIXIE SENESAC2,3,MATTHEW F. W ASSON1,4,PAULO E. LLAMBÂõAS1,VALENTINA FERRETTI1 AND PATRICK J. SULLIVAN5 1Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, NY 14853 2Cornell Laboratory of Ornithology, 159 Sapsucker Woods Road, Ithaca, NY 14850 3Animal Health Diagnostic Laboratory, College of Veterinary Medicine, Cornell University, Ithaca, NY 14853 4Appalachian Voices, 703 W. King St., Boone, NC 28607 5Department of Natural Resources, Cornell University, Ithaca, NY 14853 Abstract. To study the patterns and determinants of philopatry and breeding dispersal in the Tree Swallow (Tachycineta bicolor) we analyzed the records of 356 males and 1459 females captured in more than one breeding year around Ithaca, New York. Of these cap- tures, only 4% of male and 14% of female breeders dispersed to a new site for breeding. With our combination of intensive study areas in Tompkins County, New York, and the efforts of volunteer banders throughout New York and surrounding states, we could have detected dispersal in excess of 400 km from the initial breeding site. Randomization tests revealed, however, that breeders dispersed much shorter distances than they could have been detected. Detailed analyses of recaptures in Tompkins County showed that over a 22-km range of distances, the chances of dispersal to a new breeding site declined with the distance from the original breeding site. Females that failed to ¯edge any offspring were much more likely to disperse than females that reproduced successfully, and the probability of dispersal declined gradually with female age. The spatial scale in which swallows gather and process information appears to be much larger than for passerines that defend all-purpose territories. Key words: adult mortality, breeding success, dispersal, philopatry, spatial scale, Tach- ycineta bicolor, Tree Swallow. DispersioÂn Reproductiva y FilopatrõÂa en Tachycineta bicolor Resumen. Para estudiar los patrones y los determinantes de la ®lopatrõÂa y la dispersioÂn reproductiva en Tachycineta bicolor analizamos los registros de 356 machos y 1459 hembras capturados en maÂs de un anÄo reproductivo en los alrededores de Ithaca, New York. De estas capturas, soÂlo el 4% de los machos y el 14% de las hembras reproductivas se dispersaron a un nuevo sitio de crõÂa. Con nuestra combinacioÂn de aÂreas de estudio intensivas en el Condado de Tompkins, New York, y los esfuerzos de colaboradores voluntarios que anillaron aves a lo largo de New York y los estados circundantes, pudimos haber detectado eventos de dispersioÂnamaÂs de 400 km desde el sitio de crõÂa inicial. ExaÂmenes aleatorizados reve- laron, sin embargo, que las aves reproductivas se dispersaron a distancias mucho maÂs cortas que las que se podrõÂan haber detectado. AnaÂlisis detallados de recapturas en el Condado de Tompkins mostraron que en un rango de distancias de 22 km, las probabilidades de disper- sioÂn a un nuevo sitio de crõÂa disminuyeron con la distancia desde el sitio de crõÂa original. Las hembras que fracasaron en la crõÂa de pichones presentaron una probabilidad de disper- sarse mucho mayor que las hembras que se reprodujeron exitosamente, y la probabilidad de dispersioÂn disminuyo gradualmente con la edad de la hembra. La escala espacial a la que T. bicolor recoge y procesa informacioÂn parece ser mucho maÂs grande que la de aves paserinas que de®enden territorios de uso muÂltiple. INTRODUCTION dispersal is one of the most important life-his- All organisms are confronted by variable envi- tory responses to unavoidable habitat heteroge- ronments. For those organisms that can move, neity. Birds have some of the best-developed abilities to move, and the dispersal biology of birds is gradually becoming understood (Paradis Manuscript received 11 December 2003; accepted et al. 1998, Koenig et al. 2000, Powell and 11 June 2004. Frasch 2000, Hansson, Bensch, and Hasselquist 6 E-mail: [email protected] 2002, Paradis et al. 2002), despite the formida- [768] TREE SWALLOW BREEDING DISPERSAL 769 ble challenges of following their movements (Koenig et al. 1996, Koenig et al. 2000). Ornithologists de®ne dispersal as movement to a new breeding site from a natal or previous breeding site and distinguish it from the migra- tions that take birds to and from often distant wintering areas. Natal dispersal involves a bird's movement to its ®rst breeding site. By de®nition, it occurs only once in a bird's life, and tends to be of larger spatial scale than the movements between breeding sites, or ``breeding dispersal'' (Greenwood and Harvey 1982, Harvey et al. 1984, Lebreton et al. 2003; Winkler, in press). In multibrooded species, breeding dispersal can occur more than once in a single breeding sea- son, and all species can undertake breeding dis- persal after failed nesting attempts or between breeding seasons. It is unclear whether the un- FIGURE 1. Locations of Tree Swallow study sites derlying behavior and biology of natal and on Cornell University land to the east of Cayuga Lake (shaded) in Tompkins County, New York. Numerals breeding dispersal are really fundamentally dif- represent the unit designations for each site. Tompkins ferent. But surviving adults of breeding age County is outlined on the inset map. make repeated choices of breeding sites, increas- ing the opportunities for biologists to understand the kinds of information that are important and distances are thus not constrained by any limi- the ways that information is used in choosing tation of patchy habitat availability or vagility, sites. and this expansive potential for dispersal creates The determinants of site choice that have been a challenge to the researcher that is not present investigated include declining nest quality with in resident species. Tree Swallows possess a dis- increasing nest age (Gowaty and Plissner 1997, tinctive counteracting advantage: they are sec- Mazgajski 2003, Stanback and Rockwell 2003), ondary cavity nesters that rely on woodpeckers the sex of the bird (Harvey et al. 1978, Drilling (or humans) to create the tree holes (or nest box- and Thompson 1988, Bensch and Hasselquist es) in which they nest. Most Tree Swallows in 1991), and the success of prior nesting, either of central and western New York nest in boxes the dispersing bird (von Haartman 1949, Harvey erected for Eastern Bluebirds (Sialia sialis), and et al. 1978, Gavin and Bollinger 1988, Gowaty this habit greatly simpli®es the researcher's tasks and Plissner 1997) or its prospective neighbors of ®nding nests, protecting them from predators, (Doligez et al. 2002). Another important in¯u- and characterizing the distribution of potential ence on breeding dispersal has been mate reten- habitat. Finally, Tree Swallows are uncommonly tion or abandonment (Harvey et al. 1978, resistant to disturbance, making them easy to Bensch and Hasselquist 1991, Beheler et al. trap during nesting. 2003). Here, we concentrate on describing the Our studies of Tree Swallows around Ithaca movement patterns and exploring the nonsocial were begun with the erection of 105 nest boxes determinants of breeding dispersal in a popula- in 1985 at Cornell University's Experimental tion of Tree Swallows (Tachycineta bicolor) Ponds Unit 1 (428309N, 768289W), about 10 km nesting near Ithaca, New York. north of the Ithaca campus. Boxes were estab- lished at Experimental Ponds Unit 2 (128 boxes) METHODS in 1989, in Unit 3 along roads north of the ex- Tree Swallows are excellent subjects for study- perimental ponds (95 boxes) in 1991, on Cornell ing breeding dispersal. They ¯y every year be- farmland on Mt. Pleasant (Unit 4, 60 boxes) in tween breeding grounds throughout North 1991, along Hanshaw Road (Unit 5, 22 boxes) America and wintering areas in the Gulf Coast in 1993, and at Cornell's Harford Animal Sci- of North America, the Caribbean, and Central ence Complex (Unit 6, 131 boxes) in 2001 (Fig. America (Robertson et al. 1992). Their dispersal 1). Because of problems with House Sparrow 770 DAVID W. WINKLER ET AL. (Passer domesticus) predation, all boxes were duce an estimate of the median and range of the removed from Unit 3 after the 1993 nesting sea- expected dispersal-distance distribution for all son. Boxes at each unit are 20 m from the near- dispersals events. est neighboring box. All nest boxes on the units The uniform null model assumed that poten- were cleaned out before each breeding season, tial dispersers were equally able to ®nd and so the present study gathered no data on the ef- reach all the nesting opportunities in our entire fect of nest age on breeding dispersal (Gowaty 400-km-radius study circle. One alternative to and Plissner 1997, Mazgajski 2003). Tree Swal- this null model is that the birds searched for al- lows in the northeastern U.S. are single brooded ternative nesting sites starting at their previous (Robertson et al. 1992), and the breeding move- site and working outward from there until they ments described here are all movements between found an unoccupied site. Such local searches the breeding locations of one year and the next. produce a geometric decline in frequency with We also did not explore ®ne-scaled movements distance (e.g., Murray 1967, Waser 1985), and that birds may have made within units where we created a similar exponential null distribution territorial boundaries were vague. The move- by regressing the overall observed log probabil- ments we describe here are thus unequivocally ities of capture on distance and using the slope breeding dispersal, but ®ne-scale movements and intercept of this regression to parameterize may be worthy of further exploration in the fu- the null distribution. This null model thus takes ture. into account the reduction in nest-site availabil- Analyses are based on the sample of breeding ity and detectability with distance and mimics adults captured during 1990±2002, and thus what an animal picking a site at random starting cover potential breeding dispersal over 11 years.