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Fledgling Survival Increases with Development Time and Adult Survival Across North and South Temperate Zones

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Fledgling Survival Increases with Development Time and Adult Survival Across North and South Temperate Zones Ibis (2016), 158, 135–143 Fledgling survival increases with development time and adult survival across north and south temperate zones PENN LLOYD1,2* & THOMAS E. MARTIN3 1Percy FitzPatrick Institute, DST/NRF Centre of Excellence, University of Cape Town, Private Bag X3, Rondebosch 7701, South Africa 2Biodiversity Assessment and Management Pty Ltd, PO Box 1376, Cleveland, Qld 4163, Australia 3U.S. Geological Survey Montana Cooperative Wildlife Research Unit, University of Montana, Missoula, MT 59812, USA Slow life histories are characterized by high adult survival and few offspring, which are thought to allow increased investment per offspring to increase juvenile survival. Consis- tent with this pattern, south temperate zone birds are commonly longer-lived and have fewer young than north temperate zone species. However, comparative analyses of juve- nile survival, including during the first few weeks of the post-fledging period when most juvenile mortality occurs, are largely lacking. We combined our measurements of fledg- ling survival for eight passerines in South Africa with estimates from published studies of 57 north and south temperate zone songbird species to test three predictions: (1) fledg- ling survival increases with length of development time in the nest; (2) fledgling survival increases with adult survival and reduced brood size controlled for development time; and (3) south temperate zone species, with their higher adult survival and smaller brood sizes, exhibit higher fledgling survival than north temperate zone species controlled for development time. We found that fledgling survival was higher among south temperate zone species and generally increased with development time and adult survival within and between latitudinal regions. Clutch size did not explain additional variation, but was confounded with adult survival. Given the importance of age-specific mortality to life history evolution, understanding the causes of these geographical patterns of mortality is important. Keywords: adult survival, evolution, fledgling survival, life history. Globally, avian life-history strategies vary with lati- north temperate zone (Martin 1996). Theory sug- tude; in comparison with north temperate zone gests that low adult mortality in the southern tem- species, tropical birds have life histories typified by perate zone should favour increased parental smaller clutch sizes, longer developmental periods, investment per offspring to increase juvenile sur- higher nest predation, extended post-fledging par- vival (Skutch 1949, MacArthur & Wilson 1967, ental care and higher adult survival (Skutch 1985, Pianka 1970, Martin 2015). Increased investment Martin 1996, 2002, Ghalambor & Martin 2001, per offspring may be expressed in larger eggs Russell et al. 2004, Lloyd et al. 2014). These slow (Martin 2008), higher feeding rates per offspring life-history strategies appear to extend to the south (Martin et al. 2011, Gill & Haggerty 2012, Martin temperate zone, where adult mortality and life- 2015) or longer parental care after fledging (Rus- history expression seem to be more similar to sell et al. 2004). Increased investment and parental those of tropical species than to species of the care after fledging is expected to increase fledgling survival (Russell et al. 2004, Styrsky et al. 2005, Tarwater & Brawn 2010, Martin 2015), and varia- *Corresponding author. tion in fledgling survival can exert strong selection Email: [email protected] © 2015 British Ornithologists’ Union 136 P. Lloyd & T. E. Martin on life-history strategies (Martin 2014, 2015). temperate zones, (2) that fledgling survival Thus, low adult mortality may favour increased increases with adult survival and reduced brood parental investment in fewer young, resulting in size controlled for development time within and lower fledgling mortality (Martin 2015). As a between temperate zones, and (3) that south tem- result, we expect positive covariation of adult and perate zone species with their higher adult survival fledgling mortality rates among species, and latitu- and smaller brood sizes exhibit higher or similar dinal differences in fledgling mortality that parallel fledgling survival to north temperate zone species those observed for adults. So far, these possibilities when controlling for development time. We test have not been tested. these predictions by combining measurements of Survival of offspring after leaving the nest (i.e. fledgling survival for eight south temperate zone fledglings) can be influenced by the length of time passerines with data from published studies of spent developing in the nest, at least in the north fledgling survival from around the world. We temperate zone (Cox et al. 2014, Martin 2014). In define fledgling survival as the survival of young particular, species with higher nest predation risk through the first few weeks after leaving the nest. spend less time developing in the nest, causing their young to fledge with smaller relative body METHODS mass and reduced wing development (Cheng & Martin 2012, Martin 2014, 2015). Of critical rele- South Africa study vance, increased parental investment in species with slower life histories (i.e. species with higher We studied fledgling survival as part of an inten- adult survival and smaller clutch size) results in sive study of life-history variation (Martin et al. fledglings with better-developed wings but similar 2006, 2007, 2011) in a community of birds inhab- relative body mass compared with species with fas- iting the 2900-ha Koeberg Nature Reserve ter life histories for the same development time (33°410S, 18°260E, elevation 10 m), on the south- (Martin 2015). Body mass can be important to west coast of South Africa. The vegetation is fledgling survival in both north temperate coastal shrubland, with an average shrub height of (Krementz et al. 1989, Linden et al. 1992, Both 1–2 m (Nalwanga et al. 2004). The region has a et al. 1999) and tropical/south temperate zones Mediterranean climate with hot, dry summers and (Green & Cockburn 2001, Lloyd et al. 2009, cool, wet winters: 80% of mean annual rainfall falls Tarwater et al. 2011). However, most studies find during April to September. The breeding season of predation is the primary source of fledgling mortal- birds in the community lasts from late July to ity (Martin 2014). Consequently, traits such as early November. From 2000 to 2007 we under- wing development that can affect the ability to took intensive monitoring of nests (Martin et al. escape predators (Dial et al. 2006) may then be 2007, 2011) combined with ringing of breeding particularly important to fledgling survival. The adults with a unique combination of three colour similar body mass but greater wing development rings and a numbered metal ring. Approximately in species with slower life histories yields contrast- 1400 adults attending nests had been colour-ringed ing predictions for fledgling survival. In particular, by the end of 2007 (Lloyd et al. 2014). Nests if relative body mass is critical to fledgling survival, were located using parental behaviour, usually dur- then fledgling survival should not differ between ing the building stage, and checked at 1- to 4-day regions with differences in life histories when con- (mostly 1- to 2-day) intervals to determine clutch trolling for development time. In contrast, if size, stage transition dates and fate (Martin et al. mobility from wing development is critical, then 2006). To prevent premature fledging from the fledging survival should be greater among species nest, nestlings were measured and ringed within with slower life histories. Ultimately, the impor- 1–2 days of their primary feathers breaking pin, tance of offspring development time, and possible which occurs approximately 4–5 days before fledg- differences between slow vs. fast life histories (i.e. ing. For identification of fledglings out of the nest, adult survival, offspring number) for fledgling sur- nestlings were ringed with a numbered metal ring vival across latitudinal regions needs testing. and either: (1) a single coloured plastic ring, with We test three predictions: (1) that fledgling sur- each brood member receiving a different colour, vival increases with length of development time in or (2) a unique combination of three colour rings the nest within and between north and south (one species only). © 2015 British Ornithologists’ Union Fledgling survival comparison across latitudes 137 We determined whether young had fledged for 60 species (Supporting Information Table S1). from a nest by monitoring nests close to fledging Among the 37 north temperate zone species, at 1- to 2-day intervals and confirming that par- fledgling survival estimates included the use of ents were carrying food to young out of the nest radio-telemetry in 21 species and re-sighting of when the nest was first found empty. Thereafter, colour-ringed birds in the remainder. Among the the survival of young was monitored at weekly latter 16 species, nine species were year-round res- intervals until 21 days post-fledging, the most com- idents, six species were confidently monitored dur- mon interval among studies of fledgling survival ing the period prior to any dispersal from the natal (Martin 2014). All species monitored for fledgling territory taking place and one species (Purple Mar- survival were year-round territorial residents. As tin Progne subis) was monitored at predictable family groups did not disperse from the natal terri- roosting colonies. The survival estimates for all 28 tory post-fledging, reliance on radiotelemetry
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