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Apparent Annual Survival Estimates of Tropical Songbirds Better Reflect Life

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Apparent Annual Survival Estimates of Tropical Songbirds Better Reflect Life Received: 9 December 2016 | Revised: 23 August 2017 | Accepted: 30 August 2017 DOI: 10.1111/geb.12661 RESEARCH PAPER Apparent annual survival estimates of tropical songbirds betterreflectlifehistoryvariationwhenbasedonintensive field methods Thomas E. Martin1 | Margaret M. Riordan2 | Rimi Repin3 | James C. Mouton2 | William M. Blake2 1U.S. Geological Survey, Montana Cooperative Wildlife Research Unit, Abstract University of Montana, Missoula, Montana Aim: Adult survival is central to theories explaining latitudinal gradients in life history strategies. 2 Montana Cooperative Wildlife Research Life history theory predicts higher adult survival in tropical than north temperate regions given Unit, University of Montana, Missoula, Montana lower fecundity and parental effort. Early studies were consistent with this prediction, but 3Sabah Parks, Research and Education, Kota standard-effort netting studies in recent decades suggested that apparent survival rates in temper- Kinabalu Sabah, Malaysia ate and tropical regions strongly overlap. Such results do not fit with life history theory. Targeted marking and resighting of breeding adults yielded higher survival estimates in the tropics, but this Correspondence approach is thought to overestimate survival because it does not sample social and age classes Thomas E. Martin, Montana Cooperative Wildlife Research Unit, University of with lower survival. We compared the effect of field methods on tropical survival estimates and Montana, Missoula, MT 59812, U.S.A. their relationships with life history traits. Email: [email protected] Location: Sabah, Malaysian Borneo. Present address William M. Blake, MPG Ranch, Missoula, Time period: 2008–2016. MT 59801, U.S.A. Major taxon: Passeriformes. Funding information National Science Foundation, Grant/Award Methods: We used standard-effort netting and resighted individuals of all social and age classes of Number: DEB-1241041, DEB-1651283, 18 tropical songbird species over 8 years. We compared apparent survival estimates between IOS-1656120 these two field methods with differing analytical approaches. Editor: Erica Fleishman Results: Estimated detection and apparent survival probabilities from standard-effort netting were similar to those from other tropical studies that used standard-effort netting. Resighting data veri- fied that a high proportion of individuals that were never recaptured in standard-effort netting remained in the study area, and many were observed breeding. Across all analytical approaches, addition of resighting yielded substantially higher survival estimates than did standard-effort net- ting alone. These apparent survival estimates were higher than for temperate zone species, consistent with latitudinal differences in life histories. Moreover, apparent survival estimates from addition of resighting, but not from standard-effort netting alone, were correlated with parental effort as measured by egg temperature across species. Main conclusions: Inclusion of resighting showed that standard-effort netting alone can negatively bias apparent survival estimates and obscure life history relationships across latitudes and among tropical species. KEYWORDS apparent survival, egg temperature, embryonic development, latitudinal variation, life history, lon- gevity, resighting 1386 | VC 2017 John Wiley & Sons Ltd wileyonlinelibrary.com/journal/geb Global Ecol Biogeogr. 2017;26:1386–1397. MARTIN ET AL. | 1387 1 | INTRODUCTION produced similar results to those of Karr et al. (1990), although some studies found somewhat higher rates of apparent adult survival (Table Adult survival is a major component of demography and lifetime repro- 1). Standard-effort netting was later recognized to sample transient and ductive success, and figures prominently in life history theory young individuals that disperse or have higher mortality (Pradel, Hines, (Charlesworth, 1994; Martin, 2002, 2015; Michod, 1979; Roff, 2002; Lebreton, & Nichols, 1997). Subsequently, a time-since-marking Williams, 1966). Life history theory predicts that higher adult survival is approach was used to allow separation of apparent survival estimates in associated with slower life history strategies that include lower the year after first capture versus all subsequent years, under the fecundity and parental effort across taxa (Charnov & Krebs, 1974; assumption that transient and young individuals would disperse or die in Ghalambor & Martin, 2001; Pianka, 1970; Roff, 2002; Williams, 1966). that first year (Johnston, White, Peach, & Gregory, 1997; Pradel et al., Higher survival and slower strategies were long thought to be preva- 1997). Apparent survival estimates increased when such models were lent in the tropics (Dobzhansky, 1950; MacArthur, 1972; Pianka, 1970). used, but not substantially (e.g., Blake & Loiselle, 2013; Brawn, Karr, The perception of life history differences between the tropics and Nichols, & Robinson, 1999; Francis, Terborgh, & Fitzpatrick, 1999; John- north temperate regions was facilitated by early work on birds (e.g., ston et al., 1997). Ultimately, the apparent survival rates across seven Fogden, 1972; Lack, 1954; Snow, 1962; Snow & Lill, 1974; Willis, standard-effort netting studies of 90 total species that used time-since- 1974). However, Karr, Nichols, Klimkiewicz, and Brawn (1990) sug- marking models averaged .62, which was moderately higher but often gested that tropical birds do not have substantially higher apparent sur- overlapped survival estimates of north temperate species (Table 1). vival rates than temperate species. Given the widely documented A perception of substantially higher apparent survival in the tropics lower fecundity and parental effort in tropical species (i.e., Jetz, Seker- (i.e., .74–.89) was based on early studies that resighted colour-marked cioglu, & Bohning-Gaese,€ 2008; Martin, 2015; Martin, Martin, Olson, birds (e.g., Fogden, 1972; Snow, 1962; Willis, 1974). A review of single- Heidinger, & Fontaine, 2000; Martin, Oteyza, Boyce, Lloyd, & Ton, species studies that used colour-banding and resighting indicated that 2015; Martin et al., 2006; Moreau, 1944; Skutch, 1949, 1985; Snow, apparent survival estimates were higher than those from standard- 1962), an absence of higher survival calls into question a central tenet effort netting studies (Sandercock, Beissinger, Stoleson, Melland, & of life history theory. Hughes, 2000). Yet this interpretation was complicated because com- The apparent survival rates for tropical birds estimated by Karr et al. parison of the two approaches was based on different species in differ- (1990) were based on 10 years of capture–recapture data collected with ent environments. In addition, higher apparent survival estimates from standard-effort netting in Panama and rigorous estimation methods that previous resighting studies may be biased because they often focus on accounted for detection probability (i.e., Pollock, Nichols, Brownie, & marking territorial breeding individuals and exclude social and age Hines, 1990). These methods have been used in other locations and classes that may have lower apparent survival and higher dispersal TABLE 1 Studies of multiple songbird species that encompassed 5 years and estimated annual apparent survival probability with a Cormack–Jolly–Seber model Used annual Used (A) or actual standard- Used a netting (N) Number Annual apparent effort time-since- Used intervals for Number Location of species survival probability netting marking model resighting estimates of years Reference (a) Tropical Panama 25 .558 (.022) Y N N A 10 Karr et al. (1990) Costa Rica 11 .56 (.030) Y N N A 9 Blake and Loiselle (2002, 2008) Panama 11 .578 (.028) Y Y N A 20 Brawn et al. (1999) Brazil 31 .586 (.023) Y Y N A 4–10 Wolfe, Stouffer, and Seeholzer (2014) Ecuador 37 .596 (.016) Y Y N N 12 Blake and Loiselle (2013) Ecuador 5 .616 (.096) Y Y N A 7 Parker et al. (2006) Mexico 6 .617 (.048) Y Y N A 16 Ruiz-Gutierrez et al. (2012) Trinidad 17 .653 (.024) Y Y N A 9 Johnston et al. (1997) Peru 14 .676 (.024) Y Y N A 10 Francis et al. (1999) Puerto Rico 9 .680 (.033) Y N N A 18 Faaborg and Arendt (1995) Malaysia 14 .755 (.021) Y Y Y A 5 Martin et al. (2015) Venezuela 18 .764 (.025) Y Y Y A 7 Martin et al. (2015) (b) North temperate Arizona, USA 16 .530 (.014) Y Y Y A 21 Martin et al. (2015) Maryland, USA 8a .520 (.032)b N N N A 8 Karr et al. (1990) North America 75a .542 (.011)c N N Y A Martin (1995) N 5 no; Y 5 yes. aWoodpeckers excluded. bBirds were fed supplemental food for 6 months of the year and captured weekly for 3 months at or near feeding stations. cApparent survival rates derived from the literature and based on return rates (simple percentage surviving) instead of a Cormack– Jolly–Seber model. 1388 | MARTIN ET AL. (Francis et al., 1999; Johnston et al., 1997; Sandercock et al., 2000). At estimates from standard-effort netting alone differed from estimates the same time, birds may learn to avoid nets in the tropics, where based on the addition of resighting in the same populations. We further many remain on permanent territories and possess long-term knowl- examined whether estimates differed among analytical approaches that edge of their territories. Resighting studies provide an opportunity to included annual versus actual capture intervals and inclusion versus identify whether individuals that are never recaptured actually remain exclusion of first-year birds under time-since-marking and Barker on the study area. Yet whether resighting applied to all social and age (1997)
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