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Cavity Roosting, Philopatry, and Cooperative Breeding in the Green Woodhoopoe May Reflect a Physiological Trait

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Cavity Roosting, Philopatry, and Cooperative Breeding in the Green Woodhoopoe May Reflect a Physiological Trait CAVITY ROOSTING, PHILOPATRY, AND COOPERATIVE BREEDING IN THE GREEN WOODHOOPOE MAY REFLECT A PHYSIOLOGICAL TRAIT J. DAVID LIGON, • CYNTHIA CAREY,2 AND SANDRA H. LIGON• •Departmentof Biology,University of New Mexico,Albuquerque, New Mexico87131 USA, and 2Departmentof EPO Biology,University of Colorado,Boulder, Colorado 80309 USA ABSTRACT.--Cooperativebreeding in birds generally is related to a critical resource.A critical limiting factor for the Green Woodhoopoe(Phoeniculus purpureus) is the availability of roost cavities.Although predation at roost holes is the major sourceof mortality, wood- hoopoesinvariably roost in cavitiesor under loosebark. This dependenceon cavitiesappears to be relatedto the birds' inability to copephysiologically with low nighttime temperatures. This is the first evidenceto suggestthat a physiologicallimitation has played a major role in the evolutionof philopattyand possiblycooperative breeding in an avian species.Received 2 April 1987, accepted8 October1987. THEprevalent view of the basisof cooperative Ligon 1982), the returning woodhoopoesusu- breeding in birds is that it is related to some ally flew to a secondarycavity. In the absence kind of resource(e.g. Brown 1974, Koenig and of a backuproost hole, the birds crawled under Pitelka 1981, Emlen 1982). In some cases the loosebark to spend the night. There is no evi- putative critical resource is territorial space dencethat they ever roostedin the open. More- (Woolfenden and Fitzpatrick 1984), while in over, becausewoodhoopoes do not excavate others it may be costly self-constructedstruc- their own cavities and must compete for holes, tures suchas food storagesites (Stacey and Koe- often unsuccessfully,with numerousother cav- nig 1984) or roostcavities excavatedinto living ity-dwelling animals (J. D. Ligon and S. H. Li- trees(Ligon 1970).Although the selectivebasis gon 1982, 1988), high-quality (i.e. relatively of the nondispersalelement of avian coopera- predator-proof)roost sites are scarce.The severe tive breeding is unresolved (Staceyand Ligon interspecific competition for cavities (honey- 1987), the specificfactors prohibiting dispersal bees, rodents, other birds) means that safe cav- or, alternatively, favoring group living vary ities vary in number both spatially and tem- from speciesto species,and nondispersal of porally. Becauseof the frequent takeover of young birds is a necessaryprecondition for the cavities by other species,a key element of ter- evolution of cooperativebreeding. ritory quality is the number of potential backup We studied the behavior and ecology of the or reserve roost sites. cooperatively breeding Green Woodhoopoe Although the woodhoopoesthus require cav- (Phoeniculuspurpureus) near Lake Naivasha in ities in which to roost,this behavior appearsto the Rift Valley of Kenya (0ø40'S,36ø23'E) for be costlybecause the holes used often are in several years, 1975-1984, and found that for weak wood not chosenby other species,and thesebirds tree cavitiessuitable for roosting are the birds are vulnerable to arboreal nocturnal a critical limiting factor (J. D. Ligon and S. H. mammalian predators, such as large-spotted Ligon 1978, 1982, 1988). We followed several genets(Genetta tigrina) and possiblycats (Felis hundred birds to their roosts,usually to capture libycaand F. domesticus;Ligon and Ligon 1978). them for individual marking (386 woodhoo- Driver ants (tribe Dorylini) also capture adult poes banded). Without exception, the wood- woodhoopoesin their roost holes. Primarily as hoopoesroosted under cover. They almost al- a result of nocturnal predation at roost sites, ways entered a tree cavity and usually roosted Green Woodhoopoessuffer annual mortality in groups; as many as eight birds have been ratesabout twice as high as some other well- recorded in a single cavity (J. D. Ligon and S. studied cooperatively breeding species,about H. Ligon 1982). If, during the day, the roost 40% per year for males and 30% per year for cavity was occupied by another species,e.g. females (Ligon 1981, 1983; Ligon and Ligon honeybees(Apis mellifera; S. H. Ligon and J. D. 1988). 123 The Auk 105: 123-127. January1988 124 LIGON,CAREY, AND LIGON [Auk,Vol. 105 T^BLE1. Metabolic rates and body temperaturesof h. During this perioddried air flowed continuously Green Woodhoopoesat different ambient temper- throughthe chamberat approximately725 ml/min. atures (T•). Measurementof •rO2 was conductedin a manner Male Male Fe- similarto that describedby Dawsonand Carey(1976). I 2 male Dry air was channeledthrough flow meters,into the metabolicchambers, through tubes containing As- Body mass(g) 71.0 66.1 52.3 carite and Drierite for removal of CO2 and water va- Daytime body temperature por, respectively,and then into a Beckmanmodel 755 (øC) at Ta 27øC 42ø 42ø 40ø paramagnetic oxygen analyzer for analysis. The air Metabolic rate (ml O2.g-•.h -•) from each chamberwas sampledsequentially for 5 At T• 30.5øC 1.33 1.67 1.14 min, and room air was sampled once an hour. After At T• 27øC 1.35 1.40 1.09 steady values were obtained at the chamber air tem- At T• 19øO 1.92 2.54 1.31 perature of 30.5øC,the temperature of the constant- Body temperature (øC) at Ta temperaturebox was lowered to 27øC,and finally to 19øC, at 2400 36 ø 35 ø 33 ø 19øC.The birds were exposedto 27øCand 19øCfor at • Metabolic rates at i9øC were taken as the birds moved about. We least I h after the temperature in the chamber had were unable to obtain "resting" rates. stabilizedbefore measurementsof •O2 were made. Two or three measurements were taken at 30.5øC and 27øCand averaged to represent the value for that In view of the high risk of mortality associ- individual at those temperatures. atedwith roostingin cavities,the obviousques- Body temperatureswere taken cloacallywith a YSI tion is why woodhoopoesdo not roost among telethermometerused in conjunctionwith a YS! small- animal probe. the thorny branches and twigs of the acacia trees(Acacia xanthophloea) on the study site, as do mostother small birds,including someother RESULTS group-living speciesof similar masssuch as two Average oxygen consumption of the three speciesof Turdoidesbabblers (pets. obs.) and the birds at 30.5øCfell within 5% of the value pre- Grey-backed Fiscal Shrike (Laniusexcubitorius; dictedby the equationof Aschoffand Pohl (1970) Zack 1986). The low, 7-10øC, nighttime tem- for nonpasserinebirds measuredin the inactive peraturescharacteristic of the study site at 1,860 phase of their daily cycle (Table 1). Although m above sea level, together with the extreme we did not measure•O2 at a wideenough va- agitationshown by woodhoopoeswhen denied riety of temperaturesto describea thermoneu- accessto their roostsat dusk, suggestedthey tral zone, the similarity between the predicted might be physiologically incapable of coping and measured values and information about successfullywith low nocturnal ambient tem- typical lower critical temperatures in birds of peratures (Ligon and Ligon 1978). To address similar body mass(Calder and King 1974) sug- this possibility, we investigated thermoregu- gest that these birds were in what would be latory responsesof three captiveGreen Wood- termeda thermallyneutral condition. The hoopoesto low nocturnal ambient tempera- tures. of thesebirds remained relatively stableas am- bient temperaturewas lowered to 27øC,but in- creasedsharply when air temperaturedropped METHODS to 19øC.Although the birds were totally inac- We borrowed 2 adult male Green Woodhoopoes tive at 30.5 ø and 27øC, the elevated rates at 19øC from the Denver Zoo and 1 young female from the were coincident with frantic hopping and flut- Rio GrandeZoo in Albuquerque,New Mexico. One tering movements of the bird against the sides of the males and the female had been hatched in and roof of the metabolic chambers. These captivity;the other male was capturedfrom the wild. movements became progressively more fre- Measurementof oxygenconsumption (•O2) was quent the longer the birds remained at 19øC.To conductedusing 3.8-1metabolic chambers with black- avoid damage to the woodhoopoes,we termi- ened inner surfaces.Air temperatures within the nated the experiment at 2400 and immediately chamber were measuredwith a copper-constantan removed them from the environmental cham- thermocouple connectedto a Soltex millivolt record- er. The birdswere placedindividually in the cham- ber. The body temperature of each woodhoo- bers at 1600. The chamberswere then placed in a poe, measuredas it was taken from the chamber, constant-temperaturebox, and the birds were allowed indicatedthat despitethe increasedactivity, the to adjust to conditions in the chamber at 30.5øCfor 4 birdswere becominghypothermic (Table 1). The January1988] GreenWoodhoopoe Physiology 125 hypothermia was not an adaptive response to turnal metaboliccosts. Early selectionfor cavity low ambient temperatures; rather, it occurred roostingcould have led to an eventualloss of despite the metabolic heat production associ- thermoregulatoryabilities at low ambient tem- ated with intense locomotor activity. peratures. If cavityroosting is adaptive,low metabolic DISCUSSION rates make for more efficient use of cavities, in that 02 uptake and CO2 production in the cav- Compared with the Temperate Zone avian ities is lessthan if the birds had higher rates. speciesthat have been studied (mostly passer- This could be an important considerationin an ines), these tropical coraciiform birds demon- environment where cavities are in limited sup- strated a very different and apparently inap- ply and where several
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