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Mcdonald Et Al 2007 Bes

Mcdonald Et Al 2007 Bes

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ORIGINAL PAPER 
A critical analysis of 
 
false-feeding
 behaviorin a cooperatively breeding bird: disturbance effects,satiated nestlings or deception?
Paul G. McDonald
 &
 Anahita J. N. Kazem
 &
Jonathan Wright
Received: 27 November 2006 /Revised: 28 February 2007 /Accepted: 3 March 2007
#
Springer-Verlag 2007
Abstract
 
False feeding,
 where helpers arrive at nests withfood but fail to provision the young, has been reported inseveral cooperative species. This and other potentially
deceptive
 behavior has been interpreted as indicating that helping may operate as a signal within such socialgroups. Wecritically examine these phenomena in the provisioning behavior of the bell miner 
 Manorina melanophrys
. Exces-sively close observation distances can artificially elevate therate of false feeding in this (and other) species, but once thishad been accounted for, there was little evidence for any
deceptive
 behavior by helpers or breeders. Natural andexperimentally induced variation in the presence of a potential conspecific audience at the nest did not have anyconsistent influence upon the rate of false feeds, which waslow at 7.94% of 6,880 nest visits. Instead, encounteringunexpectedly low levels of brood demand provided a more parsimonious explanation for those visits where helpersfailed to feed nestlings or ate the food themselves. Failure tocompletely transfer a load to nestlings was more likely whenthe load contained a high proportion of sticky lerp,indicating a simple prey-transfer problem. Finally, individu-als that arrived at nests without prey were often members of neighboring breeding pairs, suggesting that these few non-feeding visits may instead involve an information-gatheringfunction. We, therefore, suggest that future studies explicitlyexclude the possibility of observer disturbance and
 all 
aspects of normal provisioning behavior before applyingthe terms
 
false feeding
 or 
 
deceptive
 and inferringanything more than straightforward helping at the nest.
Keywords
 Bellmine.White-wingedchough.Carrioncrow.Helpingatthenest .Signalinghypotheses
Introduction
Investigation of avian helping at the nest has providednumerous insights into the evolution of cooperation withinsocial groups (see Brown 1987, Emlen 1991, Koenig and Dickinson 2004, and Cockburn 2006 for reviews). The traditional view of helping as an apparently altruistic behavior has been challenged by an increasing number of studies in which helpers have been reported to
 
false feed.
That is, helpers appear to gain the benefits of being seen toattend and provision the young without incurring the costsof actually transferring food to others. Such atypical provisioning behavior has been documented in cooperativegroups of brown jays (
 Psilorhinus morio
; Lawton andGuindon 1981), bell miners (
 Manorina melanophrys
;Clarke 1984; Poiani 1993), white-throated magpie-jays (
Calocitta formosa
; Langen 1996), Arabian babblers(
Turdoides squamiceps
; Wright  1997), laughing kookabur-ras (
 Dacelo novaeguineae
; Legge 2000), and carrion crows(
Corvus c. corone
; Canestrari et al. 2004), as well as one
Behav Ecol SociobiolDOI 10.1007/s00265-007-0394-2Communicated by R. GibsonP. G. McDonaldSchool of Biological Sciences, University of Wales,Bangor, Gwynedd LL57 2UW, United KingdomP. G. McDonaldDepartment of Zoology, La Trobe University,Bundoora 3086, AustraliaA. J. N. Kazem
:
J. Wright Institute of Biology, NTNU,Trondheim 7491, NorwayP. G. McDonald (
*
)Centre for the Integrative Study of Animal Behaviour,Macquarie University,Sydney 2109, Australiae-mail: paul@galliform.bhs.mq.edu.au
 
mammalian species (meerkats,
 Suricata suricatta
; Clutton-Brock et al. 2005). However, the most frequently citedexample is that of the white-winged chough (
Corcoraxmelanorhamphos
; Boland et al. 1997). In this species,young helpers carry food to the nest area, but instead of  provisioning the nestlings, they sometimes dip the food intoa nestling
s bill without releasing it before consuming the prey item themselves. This suggests that these individualsare attempting to minimize the costs associated with providing aid, yet still be identified as a helpful individualwithin the group. False feeding is more common whenindividuals are alone at the nest, with only 3 of 89 incidentsoccurring when another individual was present (Boland et al.1997). As such, this behavior has been interpreted as a formof 
 
deception,
 sensu
 Semple and McComb (1996), wherea receiver registers a situation that is not in realityoccurring, and the signaler benefits while the receiver incurs a cost of this misinformation. Boland et al. (1997)suggest that direct benefits could accrue to deceptivehelpers in this system via one of two hypotheses proposedto account for cooperative breeding. The first, the pay-to-stay hypothesis, predicts that helping is simply a means of securing group membership and its associated benefits(Gaston 1978; Kokko et al. 2002). Individuals that fail to help sufficiently face aggression from dominant breeders or expulsion from the group (Reye 1990; Mulder andLangmore 1993). Alternatively, helping effort has beenhypothesized to be a reliable signal of individual quality,with individuals of high quality obtaining greater access tomatings or beneficial alliances (social prestige; Zahavi1977, 1995; Wright  1999). Thus, false feeding may allow individuals to retain access to the group, or be perceived asa high-quality individual, without incurring the costs of handing over the food to the nestlings.An alternative explanation for false feeds is simply that helpersbasetheirdecisiontofeedabrooduponthedifference between their own and the brood
s nutritional state (Wright 1997; Canestrari et al. 2004; Clutton-Brock et al. 2005). Individuals that are in relatively poor condition and/or exposed to unexpectedly low brood demand during a provisioning visit may benefit by consuming prey itemsthemselves, rather than feeding them to the nestlings. Thishypothesis receives circumstantial support from the fact that it is the younger white-winged chough and breeding femalecrows that exhibit the highest rates of false feeding in thestudies cited above. Younger chough are typically in poorer condition than adults (Boland et al. 1997), and breedingfemale crows face the greatest burden of all individuals over a breeding season as sole brooders and incubators of severalclutches per season (Canestrari et al. 2004). Younger brown jays (Lawton and Guindon 1981) and
 
floater 
 white-throated magpie-jays (Langen 1996) were also the most likely to make
 
 provisioning mistakes,
 and again, these arethe classes of an individual likely to be the most nutri-tionally stressed.A critical test of this second hypothesis requires that the begging response of a brood during specific visits isnegatively correlated with the propensity that an individualwithholds food from the nestlings. Boland et al. (1997)conducted a food supplementation manipulation and inferredreduced nestling hunger as a consequence (based on thenumber of begging vocalisations
 between
 provisioningevents); however, the supplementary food was equally likelyto have reduced hunger in attendants as well as nestlings.Under this scenario, a helper using a simple provisioningrule of 
 
feed nestlings when they seem hungrier than me
would show the observed reduction in the frequency of falsefeeds, thereby, precluding the need for any explanation basedon
 
deceptive
 behavior (Canestrari et al. 2004). Likewise,while Canestrari et al. (2004) found evidence suggestive of this second alternative, neither study quantified the nestling begging levels at the critical moment when provisioners arefaced with the decision to provision the young or false feed.Furthermore, under any signaling explanation, the decisionto false feed might be expected to depend on prey quality/ size (i.e., the cost to helpers of giving the prey item to thenestlings as opposed to consuming it themselves), or the presence/absence of other individuals in the vicinity. This isnot the case if false feeds merely reflect a temporal mismatch between anticipated brood demand and current state of the provisioning individual.Another possibility is that many apparently atypical provisioning behaviors are in fact a by-product of observer-induced disturbance at the nest (Wright  1997, 1999). Despite manyofthesestudiesrelyinguponrelativelycloseobservationor filming of cooperative groups at nests, specific statisticaltests to detect any effect of observer-generated disturbanceupon individual provisioning behaviors are disappointinglyrare. Behaviors such as alarm calling and mobbing of observersareobvious,butothermoresubtlebehavioraleffectsmay pass unnoticed. For example, in their observations of Florida scrub-jays (
 Aphelocoma coerulescens
), Stallcup andWoolfenden (1978) describe helper arrival at the nest without food, as well as possessive and prolonged attention aroundthe nest by a helper that even snatched food from the beaksof other visiting adults, and high levels of nest vigilanceespecially by the breeding pair. Such behaviors are verysimilar to the behavior exhibited by other species whendisturbed by observers present close to nests (Wright  1999).For example, Wright (1997) found that dominant Arabian babblers devoted more time to remaining in the nest area at the expense of foraging and provisioning activities whenobservers sat too close to nests. Further, Lawton andGuindon (1981) report that observers were sometimesmobbed by younger nest attendants during observations
 — 
the very class of helper that displayed the highest rate of false
Behav Ecol Sociobiol
 
feeds in their study. Observer-associated disturbance, there-fore, seems one of the most likely causes of many aberrant  behaviors at the nest, especially given that both the Arabian babbler and the Florida scrub-jay populations were habituat-ed to humans through hand feeding and the observationscarried out by unconcealed observers sitting very close tonests (Wrigh 1999).In this paper, we present a critical analysis of falsefeeding and other potentially
 
deceptive
 provisioning behaviors in the bell miner, a species in which false feedshave been described previously (Clarke 1984; Poiani 1993). Bell miners form large colonies that contain a number of coteries, each of which consists of a number of breeding pairs and an attending assemblage of non-breeding helpersthat are mostly male, and provision at multiple nests withintheir coterie (Clarke 1989). While some helpers are relatedto the breeding pair they assist, many are not (Conrad et al.1998; Painter et al. 2000; e.g., 52% of helpers in this study). Circumstantial evidence points to a signalingexplanation for helping in bell miners. For example, Clarke(1989) has documented five instances in which breedingmales died or disappeared, and on each occasion, thewidowed female subsequently paired with the unrelatedmale helper that had provided the greatest assistance inraising her previous brood. A subsequent study entailingexperimental removal of breeding males revealed similar  patterns of mate choice by widowed females (Jones 1998).This suggests that one of the benefits for non-kin helpersmay be in
 
showing off 
 to the female and enhancing their chances of gaining future breeding opportunities (cf. social prestige). Unusually, bell miners also produce individuallyidentifiable
 
mew
 vocalisations during provisioning andwhen they leave the nest area (McDonald et al. 2007),meaning that individuals
 visits (and, therefore, potentiallyhelping effort) may be more easily monitored by conspe-cifics than in many species. Therefore, bell miners wouldappear one of the cooperative systems most likely toinvolve helping as a signal and, thus, also greater potentialfor 
 
deceptive
 provisioning behavior and false feeding at the nest. We tested for this by conducting detailedobservation of individual provisioning behavior on a visit- by-visit basis, in conjunction with recording levels of nestling begging, as well as both natural and experimentallyinduced variation in the presence of any potential audience(e.g., the breeding male or female) near the nest.
Materials and methods
Study sitesThe study was conducted between June 2004 and December 2005 on two bell miner colonies. The first consisted of 40
 – 
45individualsattheLaTrobeUniversityWildlifeReserve,20kmnortheast of Melbourne, Australia (37°42
S, 145°02
E). ThesecondwasnearSaintAndrews,50kmnortheastofMelbourne(37°43
S, 145°03
E), and contained 120
 – 
135 birds.Molecular analysesIndividuals within colonies were color banded, and a 70-
μ 
l blood sample was collected from the alar vein for analysis.Thissamplewasstoredin70%ethanolandthentransportedtothe Australian National University, Canberra, Australia,where birds were sexed and six-loci genotyped according tothe protocols outlined in Fridolfsson and Ellegren (1999) andPainter et al. (1997), respectively. Birds were then classed aseither a breeding female or breeding male (mean pairwiserelatedness
 r 
 between breeding pairs 0.152±0.059 SE,
 n
=23 pairs), with helpers being classified into one of six groupsaccording to their sex and relatedness to the breeding female.Relatedness was assessed using Kinship v1.2 (Goodnight and Queller  1999), which calculated the likelihood of helpers being either related (primary hypothesis,
 r 
=0.5; null hypoth-esis,
 
=0), or unrelated (primary hypothesis
 
=0, nullhypothesis of 
 r 
=0.5). We calculated the ratios of likelihoods between primary and null hypotheses based on the ratiorequired to exclude 95% of 1,000 simulated pairwisecomparisons (Goodnight and Queller  1999). Based on thetests above, helpers were determined to be either 
 
related
(males,
 
=0.424±0.031 SE,
 n
=23; females,
 
=0.530±0.041 SE,
 n
=13),
 
unrelated
 (males,
 
=
0.068±0.020 SE,
n
=103; females,
 r 
=
0.170.052 SE,
 n
=17) or, if neither significantly related nor unrelated, as
 
unresolved
 (males,
=0.210.024 SE,
 n
=46; females,
 
=0.160.059 SE,
n
=8). Values of 
 
 can range between
 
1 and +1 inKinship, with negative values indicating individuals that share fewer alleles than average for the population (Queller and Goodnight  1989). Theoretically, in a randomly mating population,
 
 should approximate 0.5 for full siblings and0.25 for half-sibs and so on (Queller and Goodnight  1989).Therefore, the estimated
 
 values for our different classesof helpers are reassuringly close to those expected. For simplicity, all results are presented using relatednesscalculated relative to the breeding female; the results didnot differ if relatedness groups were assigned relative toeither (1) breeding males or (2) the mean of the breeding pair (results not presented).Monitoring of nesting attemptsOnce found, nest contents were monitored every secondday to determine the hatching date. Only one female participated in nest construction, incubation, and brooding,thereby, allowing the breeding female to be identified for each nest. To identify breeding males (genetic data were
Behav Ecol Sociobiol

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