Reproductive Success, Growth and Survival of Black-Crowned Night-Heron (Nycticorax Nycticorax) and Snowy Egret (Egretta Thula) Chicks in Coastal Virginia

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Reproductive Success, Growth and Survival of Black-Crowned Night-Heron (Nycticorax Nycticorax) and Snowy Egret (Egretta Thula) Chicks in Coastal Virginia The Auk 113(1):119-130, 1996 REPRODUCTIVE SUCCESS, GROWTH AND SURVIVAL OF BLACK-CROWNED NIGHT-HERON (NYCTICORAX NYCTICORAX) AND SNOWY EGRET (EGRETTA THULA) CHICKS IN COASTAL VIRGINIA R. MICHAEL ERWIN, JOHN G. HAIG, DANIEL B. $TOTTS,AND JEFF$. HATFIELD NationalBiological Service, Patuxent Environmental Science Center, 11410American Holly Drive, Laurel,Maryland 20708, USA ABSTRACT.--Westudied reproductive success, growth, and survivalof Black-crownedNight- Heron (Nycticoraxnycticorax) and Snowy Egret (Egrettathula) chicksin two mixed-species heronries on marsh islands in ChincoteagueBay, AccomackCounty, Virginia in 1992 and 1993. We attachedradio transmitterswith mortality sensorsto the oldest chicks (A-chicks) in 11 to 22 nestsof both speciesto monitor survival during the mid- to late nestlingperiod and into the postnestingdispersal period. For both species,we found significantdifferences between 1992and 1993in growth ratesand survival. Massgrowth ratesof chickswere higher in 1993 than in 1992for both species.Culmen-length growth ratesvaried significantlydue to year-colonyeffects for night-herons,but only for hatching order for egrets.Differences in survival rates due to hatching order were found for the egrets in both years, but were found only in 1992for night-herons.As with massgrowth rates, survival of chickswas higher in 1993 than 1992.Survival of radio-markedA-chicks did not differ between speciesor years for the period from hatchingto fledging or from fledging through the end of the study (ca. two monthspostfledging). Survival ranged from 0.80 to 1.00from the time radio transmitters were attached(ca. two weeksof age) until dispersalage (53-55 daysfor egrets;55-60 days for night-herons).After birds left the colony,survival rateswere lower during the next 40 to 55 days,ranging from 0.25 to 0.60. Theseresults suggest that, at leastfor Snowy Egrets, A-chicks may be buffered from annual variations in food conditions,but that growth and survival of other brood membersmay provide a barometerof local conditions.Despite prob- lems of variability in measurement,some of theseparameters show promiseas bioindicators of estuarineconditions. We recommendthat a cost-efficientwading-bird monitoring program would include:(1) estimatingnumbers of nestingbirds of selectedspecies (e.g. Snowy Egrets) at particular estuaries,and (2) monitoring "initial brood sizes"and survival of young to at least two weeks of age in a sample of nests for each focal species.However, additional biomonitoring (e.g. marsh forage fish, contaminantloads) is necessaryto evaluatehow well top trophic-levelorganisms such as fish-eatingbirds respondto changesin estuarinepro- duction or quality. Received12 December1994, accepted 18 March 1995. RELIABLEBIOLOGICAL INDICATORS are being indicators, we investigated the reproductive soughtby a number of federal and stateagen- success,growth and survival of young Black- ciesto monitor estuarineecosystem health and crowned Night-Herons (Nycticoraxnycticorax) integrity (Gray 1980,Diamond and Filion 1987, and Snowy Egrets(Egretta thula) at two salt- Fox and Weseloh 1987, Peakall and Shugart marsh coloniesin coastalVirginia in 1992 and 1993).Colonially nestingwading birds (Cicon- 1993.Nesting studieshave been done on these lifotrees) have been identified as one potential speciesin other areas(e.g. Frederickand Col- bioindicatorgroup by a number of researchers lopy 1989a,b, Custerand Peterson1991, Fred- (Custer and Osborn 1977, Kushlan 1993, Par- erick et al. 1992, Parsons1994), thus providing sons1994, Custer in press,Erwin and Custer in a basis for year and geographiccomparison. press),although others (Morrison 1986,Temple These other studies, however, were not de- and Wiens 1989) have expressedreservations signedto evaluatehow survivalrelates to growth about the usefulness of birds as environmental and success,since investigators did not follow indicators. birdsthrough the entire nestingperiod (but see As part of an evaluation of wading birds as Frederick et al. 1993), nor did they attempt to 119 120 ERWINET AL. [Auk,Vol. 113 cleaner and on body down with small amounts of brightly coloredfingernail polish for individual iden- tification.Aluminum leg bandsand radiotransmitters were attached(see section on late-nesting-periodsur- vival) when the birds' tarsi were of sufficient size (8 to 14 days)to retain an adult-sizedband. We assigned hatchingorders to chicksbased on observedhatching dates or, for unknown dates, we used relative size (length of the bird's culmen; Custer and Peterson 1991). A-chicks were the first hatched, B-chicks the secondhatched, and so on, through E-chicks.Chicks were followed only if they were seen alive at least ortce. Nestsuccess.--To sample nests in closeto a random manner, we selectedstudy nests from 6 to 10 areas of the colonieseach year. We then comparedthe av- erageclutch size and average"initial broodsize" (i.e. number hatched) for each speciesby year. We used Fisher's exact test (Lehmann 1975) with Monte-Carlo Fig. 1. Location of nesting coloniesin Chinco- P-valueestimate statistics (Cytel 1991)to test for dif- teagueBay, Accomack County, Virginia. ferencesbetween years. The traditionalMayfield method for estimatingnest determine survival rates beyond the fledging success(Mayfield 1975, Hensler and Nichols 1981) period. This is a major gap in the demography was not appropriate to use in our study. Instead, nest successwas calculatedin two ways. First, we com- of almostall migrantbird species,although some pared the number of birds per nest that survived 14 recent data exist for a few other waterbirds, days (i.e. the age up to which we were certain of including RoseateSpoonbills (Ajaia ajaja;Bjork finding all young on a visit); second,we compared and Powell 1994) and Great White Herons (Ar- the number of successful (i.e. •1 chick survived 14 deaherodias occidentalis; Powell and Bjork 1990) days)versus unsuccessful nests using Fisher'sexact in Florida. tests.Since brood size had a significanteffect on the Our objectivesfocused on the following ma- number of chickssurviving to 14 days(see Results), jor questions:(1) How closely coupled are re- we analyzedthe differencein the number of chicks productive success,growth rates, and survival surviving to 14 days for nestsin which at least one chick hatched for each brood size. "Initial brood size" of young for the two speciesof wading birds? was defined as the total number of chicks known to How sensitiveare they to annual changes?(2) have hatched in a nest. All chicks were included, Which speciesand nestingparameters might be regardlessof whether their hatching dates were usefuland costeffective in a biomonitoringpro- known precisely. When we found chicks whose gram? hatching(or pipping) wasnot observed,we estimated the age/hatching date by comparing culmen length STUDY AREA AND METHODS to Custerand Peterson's(1991) equation and by com- paring the chick'sinitial size to thoseof its siblings. We conductedour studiesat two large mixed-spe- Samplesizes for most brood sizeswere unequalfor cies heronries (800 to 2,000 nests) on marsh islands the two years.Only for brood sizesof three for both near Chincoteague, Accomack County, Virginia speciesin both yearswere there sufficientsamples to (37ø56'N,75ø25'W) in 1992and 1993(Fig. 1). In 1992, make meaningful comparisons;however, we ana- we restrictedour studyto the northerncolony (Cause- lyzed all brood sizes, using Fisher'sexact test with way site),but in 1993expanded the studyto include Monte-Carlo P-value estimates(Cytel 1991). Thus, the southern (Willis site) colony. Since ardeid nests comparisonswith smallnumbers of nestsmay not be are mostsusceptible to disturbancein the nest-build- very powerful. If it is assumedthat a randomsample ing to egg-layingstage (Tremblay and Ellison 1979), of brood sizes in both years was obtained,we can we limited our visitsto one early in the period (mid- combine all brood sizes and evaluate whether the May) and then one or two per week from late incu- average number of chicks produced per nest is the bation (>23 May) onward. During late incubation, samein both yearsfor both species. we marked20 to 45 egretand night-heronnests with Growth rates.--On each visit, we measured chick plasticvinyl flaggingin marshelder (Iva frutescens) mass(to nearest0.1 g) usingan Ohauselectronic bal- shrubs,and checkednests once or twiceweekly. Chicks ancethat was calibrateddaily. We alsomeasured cul- were markedon the tarsuswith piecesof coloredpipe men length (to nearest0.1 ram) from the edge of the January1996] Growthand Survival of Night-Heronand Egret Chicks 121 feathersto the tip of the culnaen.Birds whose exact ciesseparate. Then, differencescaused by year-colony hatching dateswere unknown were excludedfrom weretested for eachhatching-order combination. First, the analyses.In addition, to insure a linear relation- the two 1993 colonieswere comparedseparately, and ship,we followed Custerand Peterson's(1991) meth- then the 1992 colony was added. odologyof only includingmeasurements from birds To estimate survival of a sample of birds during between ages5 and 18 days when calculatingage- the late-nestingperiod until dispersal,we attached massrelationships, and between ages3 and 18 days small (10-g) oval radios with mortality sensorsto for calculatingage-culnaen relationships. To avoidau- A-chicks between the ages of 7 and 21 days (œ= 2 tocorrelation,we calculatedindividual chick growth weeks). Radio transmitters were attached to blank alu- slopesfor the relationshipsbetween mass and age,
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