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Babbitt, G.A. and P.C. Frederick. 2008

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Babbitt, G.A. and P.C. Frederick. 2008 Zoo Biology Phenology of body mass changes during reproduction in a nomadic, tropical waterbird, the Scarlet Ibis (Eudocimus ruber) For Peer Review Journal: Zoo Biology Manuscript ID: ZOO-07-079.R2 Wiley - Manuscript type: Research Article Date Submitted by the n/a Author: Complete List of Authors: Babbitt, Gregory; Arizona State University, The Biodesign Institute Frederick, Peter; University of florida, Wildlife Ecology and Conservation Keywords: avian, nutrition, reproduction, nomadism John Wiley & Sons Page 1 of 18 Zoo Biology 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 For Peer Review 19 20 21 Phenology of body mass changes during reproduction i n a nomadic, 22 23 tropical waterbird , the Scarlet Ibis (Eudocimus ruber ) 24 25 26 27 GREGORY A. BABBITT 1,2 AND PETER C. FREDERICK 3 28 29 1Center of Evolutionary Functional Genomics, The Biodesign Institute, 30 31 Arizona State University, Tempe, AZ 85287 -5001 32 33 2Corresponding Author: [email protected] 34 35 3Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, 36 37 FL 32605 38 39 40 41 Running head – Reproduction and Nutrition in Scarlet Ibis 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 John Wiley & Sons Zoo Biology Page 2 of 18 2 1 2 3 4 5 In birds, the pre -reproductive buildup of endogenous energy reserves (e.g. body 6 7 fat) is highly variable and is often thought to be a strategy evolving in response to either 8 seasonal and/or unpredictable changes in breeding conditions. Nomadic behavior is also 9 10 thought to be an adaptation to unpredictable resource distribution in both space and time. 11 12 Because of the difficultly in obtaining a longitudinal time series of body mass es for free - 13 14 living individuals of highly nomadic species, the relationship b etween nomadism and 15 16 endogenous energy storage has not been explored. In this study, we investigated pre - 17 reproductive energy storage in a large free -flighted captive colony of highly nomadic 18 For Peer Review 19 waterbird, the Scarlet Ibis, Eudocimus ruber . We used size -corre cted body mass as an 20 21 index of body condition both prior to and during breeding. We compared both breeders 22 23 and non -breeders body condition prior to nesting. We also prevented a sub -sample of the 24 birds from gaining mass prior to nesting and compared their nesting success with a 25 26 control group that was allowed to feed freely. While significant differences were found 27 28 in pre -reproductive body condition s of br eeders and non -breeders, we were unable to 29 30 control breeding by manipulating pre -reproductive condition, most likely because of the 31 32 ability of some birds to rapidly change body condition within several days or weeks prior 33 to nesting. We conclude that pre -reproductive energy stor age is important for nesting 34 35 success in both sexes of this highly nomadic specie s, however energy stores are highly 36 37 labile and can be rapidly obtained through pre -nesting hyperphagia. 38 39 40 KEYWORDS – avian, nutrition, reproduction, nomadism 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 John Wiley & Sons Page 3 of 18 Zoo Biology 3 1 2 3 INTRODUCTION 4 5 Breeding birds rely to varying degrees upon endogenous energy reserves , such as 6 7 stored fat for reproduction . Reliance upon such reserves (“capital breeders”) is typified 8 9 by migrants breeding at high latitude, often faced with predictable seasonal food 10 11 shortage s (Raveling 1979, Ankey and MacInnes, 1978, Drent and Daan 1980 , but see 12 13 also Meijer and Drent 1999 , Gauthier and Hobson 2003 ). Reliance on energy stores in 14 early reproduction may also represent a strategy that evolves in animals and plants in 15 16 response to breeding conditions that are unpredictable in space or time ( Jon sson 1997) . 17 18 Reliance upon storedFor fat or protein, Peer however, Reviewcarries energetic costs of flight and storage, 19 20 implying that where local resources are at least seasonally abundant, reliance upon local 21 resources (“income breeding”) may be more efficient (Jonsson 1997) . 22 23 Many energetic strategies other than the storage of fat exist by which individuals 24 25 can cope with the risk of temporal and spatial uncertainty in finding enough food to 26 27 support a reproductive effort. Pre and post -breeding life history adaptations t o temporally 28 unpredictable resources include increased life -span, prolonged or intermittent breeding 29 30 seasons (Nur and Sydeman 1999, Cam et al. 1998), variable clutch size (Mock and 31 32 Forbes 1995), facultative brood reduction (Amundsen and Slagsvold 1998, For bes and 33 34 Mock 1996), and nomadism (Brown and Hopkins 1996). Colonial breeding and local 35 36 enhancement through social foraging also represent adaptations to spatially unpredictable 37 resources (Brown and Brown 1996, Clark and Mangel 1984, Poysa 1992). Such 38 39 stra tegies may allow for less reliance upon stored reserves for breeding even when 40 41 reproduction is limited by resource distribution or availability (Meijer and Drent 1999). 42 43 It is not yet clear how to predict when and to what extent birds should be reliant up on 44 body reserves for breeding , since there seem to be multiple fitness solutions to the 45 46 problem of breeding energetics. The spatial and temporal scale of resource predictability 47 48 may be key in understanding these solutions. Resource use has been relativel y well - 49 50 studied in birds that have predictable but highly seasonal environments (high latitude 51 52 breeders, e.g. Lesser Snow Goose , Chen caerulescens ( Carey, 1996) , and in situations 53 where food is temporally predictable but spatially unpredictable (seabird s, W hittow and 54 55 Rahn 1984 , Hull et al. 2002 ). But it is less well understood for animals that face 56 57 58 59 60 John Wiley & Sons Zoo Biology Page 4 of 18 4 1 2 3 unpredictable resources at both large time and large spatial scales (eg, will food be 4 5 present at all this year or in five, and will it be here or three hundred km away?). 6 7 Here, we investigate the importance of body reserves to reproduction for a 8 9 nomadic, tropical breeder, the Scarlet Ibis ( Eudocimus ruber ) that typically may have 10 11 large uncertainties in predictability of food resources at large spatial and temporal scales . 12 Scarlet Ibises are considered by some to be conspecific with the White Ibis ( Eudocimus 13 14 albus see Ramo and Busto 1987 , H ancock et al. 1992) and a t minimum , the two are a 15 16 closely related superspecies . See the American Ornithologists Union checklist at 17 18 http://www.aou.org/checklist/index.php3For Peer. Review 19 Both Scarlet and White ibises show adaptations to unpredictable breeding and 20 21 feeding conditions, including central place foraging, colonial breeding, reg ional 22 23 nomadism, low breeding site fidelity, hatching asynchrony and brood reduction, and wide 24 25 postbreeding wandering ( Kushlan and Bildstein, 1992 , Bildstein 1993 , Frederick et al. 26 199 6, Martinez and Rodrigues 1999 ). The amount of energy White Ibises consu me 27 28 during breeding has been estimated (Kushlan 1977 ), but there is no information about the 29 30 relative use of fat reserves vs local resources during the breeding cycle. The nomadic 31 32 behavior of the White Ibis makes the long term study of marked birds nearly impossible, 33 34 therefore information relating pre -nesting energetics to reproductive success is difficult to 35 obtain. In this study, we rely upon a large breeding captive population to address basic 36 37 questions about energetic strategies used by a nomadic speci es such as this. 38 39 We report on the relationship between ibis bo dy condition and reproduction in a 40 41 large free -flighted captive colony where the birds breed regularly, where birds are 42 individually marked, where age and sex were known, and where we could c ontrol food 43 44 type and amount. We hypothesized that fat levels and body condition pr ior to breeding 45 46 may be related to the initiation and success of breeding. To test this prediction we used 47 48 two approaches. First, we attempted to manipulate body condition through a 49 50 supplemental feeding program and compared reproduction (initiation rate and success) of 51 supplemented and un supplemented birds. Second, we compared body condition indices 52 53 of successful and unsuccessful breeders. Because reproductive performance can also be 54 55 influenced by other factors such as age (Saether 1990), experience (Ratcliffe et al. 1998), 56 57 58 59 60 John Wiley & Sons Page 5 of 18 Zoo Biology 5 1 2 3 time of season (Wilson and Cooper 1998), and mate quality (Otter et al. 1999), we 4 5 monitor ed these conditions either directly or indirectly. 6 7 8 9 METHOD S 10 11 12 The Study Site 13 14 15 During the spring breeding seasons of 1998 and 1999, we studied the 16 17 reproduction of over 400 individually color -banded Scarlet Ibises in a 3 ,082m 2 mixed 18 For Peer Review 19 species aviary at Disney’s Animal Kingdom in Lake Beuna Vista , F lorida . Other speci es 20 21 housed there included Caribbean Flam ingo (Phoenicopterus ruber ), Roseate Spoonbills , 22 (Platalea ajaja ) and several species of small heron. The enclosure was nearly 20m high 23 24 at some points and included some mature trees.
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