140 ShortCommunications andCommentaries LITERATURE CITED [Auk,Vol. 114 MOCK, D. W., AND B. J. PLOGER.1987. Parental ma- CHARNOV,E. L., AND J. R. KREBS. 1974. On clutch size and fitness. Ibis 116:217-219. nipulationsof optimal hatchasynchronyin Cattle Egrets:An experimentalstudy. Animal Behaviour GIBOONS, D.W. 1987. Hatching asynchronyreduces parental investment in the Jackdaw. Journal of Animal Ecology56:403-414. LACK,D. 1947. The significanceof clutch size. Ibis 89:302-352. LACK,D. 1954. The natural regulation of animal numbers.Oxford University Press,Oxford. MAGRATH,R.D. 1990. Hatching asynchrony in altricial birds.BiologicalReviewsof the Cambridge Philosophical Society 65:587-622. MOCK,D. W., AND L. S. FORGES.1994. Life-history consequences of arian broodreduction.Auk 111: 35:150-160. REZNIK,D. 1985. Costsof reproduction: An evaluation of the empirical evidence. Oikos 44:257267. STOLESON,$. H., AND $. R. BEISSINGER. 1995. Hatch- ing asynchrony and the onset of incubation in birds,revisited:When is the criticalperiod?Current Ornithology 12:191-270. WILLIAMS, G. C. 1966. Natural selection, the costs of reproduction,and a refinement of Lack's principle. American Naturalist 100:687-690. Received 12 December 1994,accepted 6 July1995. 115-123. The Auk 114(1):140-141, 1997 Abiotic Factorsand PreroostingBehavior of Greylag Geese:A Comment STg?H•q G. REE•S • D•partementde biologie,Universit•de Moncton,Moncton,New BrunswickEIA 3E9, Canada In a paper on the preroostingbehaviorof Greylag Geese (Anser anser), Schmitt (1994) concluded that "abiotic factorsdetermine departuretime but do not disturbflockcohesion"(p. 763)and that "noneof the variationin departuretime is due to variationin social interactionsleadingto flocksynchronization" (p. 762). Here, I arguethatSchmitt(1994)hasnot convincingly shown that abiotic factors have no influence on flock cohesionduring departure to the roost, that he has not clearly shown which abiotic factorsinfluence departure time, and that he has not really shown a lack of correlationbetween departuretime and flock synchronization. Schmitt's conclusion that abiotic factors do not in- over, the power of the statisticaltestwas not given. Forbes(1990)madea convincingargumentthat conclusions basedon negativeresults(lackof statistically significanteffects)shouldbe accompaniedby power analyses. Schmittstatedthat abiotic factorsinfluence depar- ture time,a conclusionthat hasbeenreachedin many other studies(seereferencesin his paper).However, it is not clear exactly which factorswere involved in the caseof Greylag Geese.The multiple regression analysisin Schmitt'stable 1 showsthat "illumination decrease" is the main factor. But a footnote to table 1 reveals that "illumination decrease" was calculated as the "difference between values at sunset and take fluenceflockcohesionis at oddswith his results(p.760) that "synchronization was lower on rainy than on off of flock."Obviously,a parametercloselyrelated cloudy and sunny evenings (both P < 0.001), but cloudy and sunny evenings did not differ." Schmitt went on to say that rainy eveningswere colder and darker than cloudy and sunny evenings.Therefore, it is likely that temperatureand/or light intensity, if not rain itself, affected synchronization. Unfortunately,no figuresor tableswith flocksynchronization as the dependentvariable were presented.Schmitt in the calculation of "illumination to the dependent variable (departure time) was used decrease," and therefore, departure time and "illumination decrease,"from the start, were not independent from eachother. Thus,it is not surprisingthat illumination decreaseexplained as much as 92% of the variation in departure time. It is obvious that the later the departuretime,the lowerthe light intensityat that time, andthe morepositive(lessnegative)the illumination reported that a multiple regressionanalysis(on data from only one season)failed to correlateany abiotic decrease, as defined. Later in the discussion, Schmitt factorwith synchronization,but rainfall apparently was not included as an independent variable. More- quickly light intensitychanged,but there is no mention of how and at what time of day this rate was calculated.The argumentof inherent relatednessbetween independent and dependent variables also couldbe usedfor other"independent"variablesused • E-mail: [email protected] seemed to redefine illumination decrease as how January1997] ShortCommunications andCommentaries 141 in theregression, suchasillumination attakeoff, tem- views nightfall asthe time pastwhich the flock cannot perature at takeoff, and temperature decrease.Moreover, the assumptionof noncollinearity among in- fly, then the closerto nightfall the flock decidesto leave, the lesspotential there is for departure to be dependent variables (Zar 1984) probably was not stretchedin time). However, Schmitt did not explore checkedbefore the multiple regressionwas conduct- this rationale. ed (if it had been, I doubt that three variables related to illumination would have been included all at once). Perhapsthis rationale should have been given; indeed, the 1986 data in Schmitt's figure 3 seem to indicate that later departures were correlated with The analysisshould have been done only with day length, light intensityat sunset,temperatureat sunset (if unrelatedto light intensity),and synchronization of flock as independent variables. Schmitt reported that geeseleft for the roostearlier on rainy than on cloudythan on sunny days,and that temperatureand light intensitywere lower on rainy than on cloudy than on sunny days. Basedon this, he statedthat "departure time changedwith temperature and illumination level; the colder and darker the evening, the earlier the geesetook off" (p. 761). Schmitt correctly pointed out that becausethe two factorsseemto be correlated,only one of them could be involved. In the discussion,he appeared to discountthe importanceof temperaturebasedon a rather goodargumentaboutthe lack of thermoregulatory stressin summer and the results of his multiple regression.However, for reasonsexplained above, the multiple regressionanalysisshould be redone, and the results are unknown at the moment. Basedon a purported lack of correlation between departure time and flock synchronization,Schmitt claimedthat "neither early nor late takeoff resultsin low synchronization."Although I have no qualms with this statementbasedon a visual inspectionof his figure 1 (I would have said "resultsfrom" rather than "results in" to be consistentwith the position of the axesand the hypothesispresentedin the introduction), a linear correlation is not appropriate to support sucha claim. If both early and late departure times led to lower synchronization, then the data points in figure 1 would be positioned in the shape of a curve (concave side to the left), and this would call for curvilinear regression,not Pearson'sproductmoment coefficients. Linear correlation would greater flock synchronization.Following Schmitt's definitionof flocksynchronization anddeparturetime, I measuredon his figure 3 the length of the bars(flock cohesion)and the distancefrom the midpoint of the bars to the dotted line showing sunset (departure time), and then recalculated the correlation coefficients between these two variables for 1986 and 1987. For 1986,my resultswere completelydifferent from thosereportedby Schmitt.I obtainedan r of -0.539 (n = 57,P < 0.001).Surprisingly,there is no coherence between the 1986datashown on his figure 3 and those shownon his figure 1. Coherenceappearsto be better for the 1987 data, and for them I obtained an r of -0.076, (n = 41, P > 0.5). Interestingly, if only one data point (that of 15 August 1987) is eliminated, the coefficientjumpsto -0.314 (n = 40, P < 0.05).It seems that later departuresare correlatedwith greaterflock synchrony,although it remains to be ascertained which variable is causaland which is dependent.If it can be shown that departure time is causal, then Schmitt'shypothesis(p. 759)that socialfactorsrelated to synchronydo not influencedeparturetime is still valid, albeit still untestedin Greylag Geese. LITERATURE CITED FORBES, L. S. 1990. A note on statisticalpower. Auk 107:438-439. SCI-IMITT•A. 1994. Influence of abiotic factors on preroostingbehaviorof GreylagGeese(Anseranser). Auk 111:759-764. ZAR,J.H. 1984. Biostatistical analysis,2nd ed. Prentice Hall, EnglewoodCliffs, New Jersey. have been appropriate to test the idea that greater synchronizationcouldoccurduring late departure(if one ReceivedI November1994, accepted6 July 1995.
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