Null Hypothesis Testing: Problems, Prevalence, And An Alternative .

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Null Hypothesis Testing: Problems, Prevalence, and an AlternativeAuthor(s): David R. Anderson, Kenneth P. Burnham, William L. ThompsonSource: The Journal of Wildlife Management, Vol. 64, No. 4 (Oct., 2000), pp. 912-923Published by: Allen PressStable URL: http://www.jstor.org/stable/3803199Accessed: 01/02/2009 12:40Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available rms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unlessyou have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and youmay use content in the JSTOR archive only for your personal, non-commercial use.Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained herCode acg.Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printedpage of such transmission.JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with thescholarly community to preserve their work and the materials they rely upon, and to build a common research platform thatpromotes the discovery and use of these resources. For more information about JSTOR, please contact support@jstor.org.Allen Press is collaborating with JSTOR to digitize, preserve and extend access to The Journal of WildlifeManagement.http://www.jstor.org

NULLHYPOTHESISTESTING:PROBLEMS,PREVALENCE,AND ANALTERNATIVEDAVIDR. ANDERSON,'2ColoradoCooperativeFish and WildlifeResearch Unit,Room 201 CO 80523, USAKENNETHP. BURNHAM,1ColoradoCooperativeFish and WildlifeResearch Unit,Room201 CO 80523, USAWILLIAML. THOMPSON,U.S. ForestService, RockyMountainResearch Station,316 E. MyrtleSt., Boise, Idaho83702, USAAbstract: This paper presents a review and critique of statistical null hypothesis testing in ecological studiesin general, and wildlife studies in particular, and describes an alternative. Our review of Ecology and the Journalof Wildlife Management found the use of null hypothesis testing to be pervasive. The estimated number of Pvalues appearing within articles of Ecology exceeded 8,000 in 1991 and has exceeded 3,000 in each year since1984, whereas the estimated number of P-values in the Journal of Wildlife Management exceeded 8,000 in1997 and has exceeded 3,000 in each year since 1994. We estimated that 47% (SE 3.9%) of the P-valuesin the Journal of Wildlife Management lacked estimates of means or effect sizes or even the sign of thedifference in means or other parameters. We find that null hypothesis testing is uninformative when no estimates of means or effect size and their precision are given. Contrary to common dogma, tests of statisticalnull hypotheses have relatively little utility in science and are not a fundamental aspect of the scientific method.We recommend their use be reduced in favor of more informative approaches. Towards this objective, wedescribe a relatively new paradigm of data analysis based on Kullback-Leibler information. This paradigm isan extension of likelihood theory and, when used correctly, avoids many of the fundamental limitations andcommon misuses of null hypothesis testing. Information-theoretic methods focus on providing a strength ofevidence for an a priori set of alternative hypotheses, rather than a statistical test of a null hypothesis. Thisparadigm allows the following types of evidence for the alternative hypotheses: the rank of each hypothesis,expressed as a model; an estimate of the formal likelihood of each model, given the data; a measure of precisionthat incorporates model selection uncertainty; and simple methods to allow the use of the set of alternativemodels in making formal inference. We provide an example of the information-theoretic approach using dataon the effect of lead on survival in spectacled eider ducks (Somateria fischeri). Regardless of the analysisparadigm used, we strongly recommend inferences based on a priori considerations be clearly separated fromthose resulting from some form of data dredging.JOURNALOF WILDLIFEMANAGEMENT64(4):912-923Key words: AIC, Akaike weights, Ecology, information theory, Journal of Wildlife Management, KullbackLeibler information, model selection, null hypothesis, P-values, significance tests.Theoretical and applied ecologists continuallystrive for rigorous, objective approaches formaking valid inference concerning sciencequestions. The dominant, traditional approachhas been to frame the question in terms of 2contrasting statistical hypotheses: 1 representing no difference between population parameters of interest (i.e., the null hypothesis, Ho) andthe other representing either a unidirectional orbidirectional alternative (i.e., the alternative hypothesis, Ha). These hypotheses basically correspond to different models. For example,when comparing 2 groups of interest, the assumption is that they are from the same population so that the difference between their true means is 0 (i.e., Ho is I,l - (2 0, or ti 1-2).1 Employedby U.S. Geological Survey, Division ofBiologicaL Resources.2 E-mail:anderson@picea.cnr.colostate.eduA test statistic is computed from sample dataand compared to its hypothesized null distribution to assess the consistency of the data withthe null hypothesis. More extreme values of thetest statistic suggest that the sample data are notconsistent with the null hypothesis. A substantially arbitrary level (a) is often preset to serveas a cutoff (i.e., the basis for a decision) for statistically significant versus statistically nonsignificant results. This procedure has various names,including null hypothesis testing, significancetesting, and null hypothesis significance testing.In fact, this procedure is a hybridization ofFisher's (1928) significance testing and Neymanand Pearson's (1928, 1933) hypothesis testing(Gigerenzer et al. 1989, Goodman 1993, Royall1997).There are a number of problems with the application of the null hypothesis testing ap-912

J. Wildl. Manage. 64(4):20000(180160 140 -120I 10080z 60.4020-1940s 1950s 1960s 1970s 1980s 1990sDecade EEcology* Bus./EconomicsIEStatsticsMI11SocialSciences * AllMedicineFig. 1. Sample of articles,based on an extensive samplingof the literatureby decade, in variousdisciplinesthat questionedthe utilityof hown for the 1990s were extrapolatedbased onsample resultsfromvolumeyears 1990-96.proach, some of which we present herein (Carver 1978, Cohen 1994, Nester 1996). Althoughdoubts among statisticians concerning the utilityof null hypothesis testing are hardly new (Berkson 1938, 1942; Yates 1951; Cox 1958), criticisms have increased in the scientific literaturein recent years (Fig. 1). Over 300 referencesnow exist in the scientific literature that warnof the limitations of statistical null hypothesistesting. A list of citations is located at http://www. cnr. colostate. edu/-anderson/thompsonl.htmland . The former website alsoincludes a link to a list of papers supporting theuse of tests. We believe that few wildlife biologists and ecologists are aware of the debateregarding null hypothesis testing among statisticians. Discussion and debate have been particularly evident in the social sciences, where atleast 3 special features (Journal of ExperimentalEducation 61(4); Psychological Science 8(1); Research in the Schools 5(2)) and 2 edited books(Morrison and Henkel 1970, Harlow et al.1997) have debated the utility of null hypothesistests in scientific research. The ecological sciences have lagged behind other disciplines withrespect to awareness and discussion of problemsassociated with null hypothesis testing (Fig. 1;Yoccoz 1991; Cherry 1998; Johnson 1999).We present information concerning prevalence of null hypothesis testing by reviewing papers in Ecology and the Journal of WildlifeManagement. We chose Ecology because it iswidely considered to be the premier journal inthe field and hence, should be indicative of sta-HYPOTHESISTESTING* Anderson et al.913tistical usage in the ecological field as a whole.We chose the Journal of Wildlife Managementas an applied journal for comparison. We reviewtheoretical or philosophical problems with thenull hypothesis testing approach as well as itscommon misuses. We offer a practical, theoretically sound alternative to null hypothesis testing and provide an example of its use. We conclude with our views concerning data analysisand the presentation of scientific results, as wellas our recommendations for changes in editorialand review policies of biological and ecologicaljournals.PROBLEMS WITH NULL HYPOTHESISOR SIGNIFICANCETESTINGThe fundamental problem with the null hypothesis testing paradigm is not that it is wrong(it is not), but that it is uninformative in mostcases, and of relatively little use in model orvariable selection. Statistical tests of null hypotheses are logically poor (e.g., the arbitrarydeclaration of significance). Berkson (1938) wasone of the first statisticians to object to the practice.The most curious problem with null hypothesis testing, as the primary basis for data analysis and inference, is that nearly all null hypotheses are false on a priori grounds (Johnson1995). Consider the null H: o0 01 02 . 05, where 00 is an expected control responseand the others are ordered treatment responses(e.g., different nitrogen levels applied to agricultural fields). This null hypothesis is almostsurely false as stated. Even the application ofsawdust would surely make some difference inresponse. The rejection of this strawman hardlyadvances science (Savage 1957), nor does it givemeaningful insights for conservation, planning,management, or further research. These issuesshould properly focus on the estimation of effects or differences and their precision and noton testing a trivial (uninformative) null. Othergeneral examples of a priori false null hypotheses include (1) Ho: p,c jx (mean growth rateis equal in control vs. aluminum-treated bullfrog, Rana catesbeiana); (2) Ho: SjC SjD (survival probability in weekj is the same for control vs. lead-dosed gull chicks, Larus spp.), and(3) Ho: pyx 0 (zero correlation between variables Y and X). Johnson (1999) provided additional examples of null hypotheses that areclearly false before any testing was conducted;the focus of such investigations should properly

914HYPOTHESISTESTING* Anderson et al.estimate the size of effects. Statistical tests ofsuch null hypotheses, whether rejected or not,provide little information of scientific interestand, in this respect, are of little practical use inthe advancement of knowledge (Morrison andHenkel 1969).A much more well known, but ignored, issueis that a particular a-level is without theoreticalbasis and is therefore arbitrary except for theadoption of conventional values (commonly 0.1,0.05 or 0.01, but often 0.15 in stepwise variableselection procedures). Use of a fixed a-level arbitrarily classifies results into biologically meaningless categories significant and nonsignificantand is relatively uninformative. This NeymanPearson approach is an arbitrary reject or notreject decision when the substantive issue is oneof strength of evidence concerning a scientificissue (Royall 1997) or estimation of size of aneffect.Consider an example from a recent issue ofthe Wildlife Society Bulletin, "Response ratesdid not vary among areas (X2 16.2, 9 df, P 0.06)." Thus, the null must have been R1 R2 R3 . Rio; however, no estimates of theresponse rates (the RA)or their associated precision or even sample size were provided. Hadthe P-value been 0.01 lower, the conclusionwould have been that significant differenceswere found and the estimates Ri and their precision given. Alternatively, had the arbitrary alevel been 0.10 initially, the result would havebeen quite different (i.e., response rates variedamong areas, x2 16.2, 9 df, P 0.06). Here,as in most cases, the null hypothesis was falseon a priori grounds. Many examples can befound where contradictory or nonsensical results have been reported (Johnson 1999). Legalhearings concerning scientific issues are unproductive and lead to confusion when 1 partyclaims significance (based on a 0.1), whereasthe opposing party argues nonsignificance(based on a- 0.05).The cornerstone of null hypothesis testing,the P-value, has problems as an inferential toolthat stem from its very definition, its applicationin observational studies, and its interpretation(Cherry 1998, Johnson 1999). The P-value is defined as the probability of obtaining a test statistic at least as extreme as the observed one,conditional on the null hypothesis being true.There are 2 important points to consider aboutthis definition. First, a P-value is based not onlyon the observed result (the data collected), butJ. Wildl. Manage. 64(4):2000also on less likely, unobserved results (data setsnever collected) and therefore overstates theevidence against the null hypothesis (Bergerand Sellke 1987, Berger and Berry 1988). A Pvalue is more of a statement about the eventsthat never occurred than it is a concise statement of the evidence from an actual observedevent (i.e., the data). Bayesians (people makingstatistical inferences using Bayes' theorem;Gellman et al. 1995) find this property of Pvalues objectionable; they tend to avoid null hypothesis testing in their paradigm.A second consequence of its definition is thata P-value is explicitly conditional on the null hypothesis (i.e., it is computed based on the distribution of the test statistic assuming the nullhypothesis is true). The null distribution of thetest statistic (e.g., often assumed to be F, t, z,or x2) may closely match the actual samplingdistribution of that statistic in strict experiments, but this property does not hold in observational studies. In these latter studies, thedistribution of the test statistic is unknown because randomization was not done, and hencethere are problems with confounding factors(both known and unknown). In observationalstudies, the distribution of the test statistic under the null hypothesis is not deducible fromthe study design. Consequently, the form of thedistribution is not known, only naively assumed,which makes interpretation of test results problematic.It has long been known and criticized that theP-value is dependent on sample size (Berkson1938). One can always reject a null hypothesiswith a large enough sample, even if the truedifference is trivially small. This points to thedifference between statistical significance andbiological importance raised by Yoccoz (1991)and many others before and since. Anotherproblem is that using a fixed a-level (e.g., 0.1)to decide to reject or not reject the null hypothesis makes little sense as sample size increases. Here, even when the null hypothesis istrue and sample size is infinite, a Type I error(rejecting a null that is true) still occurs withprobability a (e.g., 0.1), and therefore this approach is not consistent (theoretically, a shouldgo to zero as n goes to infinity). Still anotherissue is that the P-value does not provide information about either the size or the precision ofthe estimated effect. The solution here is tomerely present the estimate of effect size and ameasure of its precision.

J. Wildl. Manage. 64(4):2000A pervasive problem in the use of P-values isin their misinterpretation as evidence for eitherthe null or alternative hypothesis (see Ellison1996 for recent examples of such misuse). Theproper interpretation of the P-value is based onthe probability of the data given the null hypothesis, not the converse. We cannot accept orprove the null hypothesis, only fail to reject it.The P-value cannot validly be taken as the probability that the null hypothesis is true, althoughthis is often the interpretation given. Similarly,the magnitude of the P-value does not indicatea proper strength of evidence for the alternativehypothesis (i.e., the probability of Ha, given thedata), but rather the degree of consistency (orinconsistency) of the data with Ho (Ellison1996). Phrases such as highly significant (oftendenoted as ** or even ***) only reinforce thiserror in interpretation of P-values (Royall 1997).Presentation of only P-values also limits theeffectiveness of (future) meta-analyses. There isa strong publication bias whereby only significant P-values tend to get reported (accepted) inthe literature (Hedges and Olkin 1985:285-290,Iyengar and Greenhouse 1988). Thus, the published literature is itself biased in favor of results arbitrarily deemed significant. It is important to present parameter estimates (effect size)and their precision from any well designedstudy, regardless of the outcome; these becomethe relevant data for a meta-analysis.A host of other problems exist in the null hypothesis testing paradigm, but we will mentiononly a few. We generally lack a rigorous theoryfor testing null hypotheses when a model contains nuisance parameters (e.g., sampling probabilities in capture-recapture studies). The distribution of the likelihood ratio test statistic between models that are not nested is unknownand this makes comprehensive analysis problematic. Given the prevalence of null hypothesistesting, we warn against the invalid notion ofpost-hoc or retrospective power analysis (Goodman and Berlin 1994, Gerard et al. 1998) andnote that this practice has become more common in recent years.The central issues here are twofold. First, scientists are fundamentally interested in estimates of the magnitude of the differences andtheir precision, the so-called effect size. Is thedifference trivial, small, medium, or large? Isthis difference biologically meaningful? This isan estimation problem. Second, one often wantsto know if the differences are large enough toHYPOTHESISTESTING Anderson et al.915justify inclusion in a model to be used for inference in more complex science settings. Thisis a model selection problem. These central issues that further our understanding and knowledge are not properly addressed with statisticalhypothesis testing. Statistical science is muchmore than merely significance testing, eventhough many statistics courses are still offeredwith an unfounded emphasis on null hypothesistesting (Schmidt 1996). Many statisticians question the practical utility of hypothesis testing(i.e., the arbitrary oa-levels, the false null hypotheses being tested, and the notion of significance) and stress the value of estimation of effect size and associated precision (Goodmanand Royall 1988, Graybill and Iyer 1994:35).PREVALENCE OF FALSE NULLHYPOTHESES AND P-VALUESWe randomly sampled 20 papers in the Articles section from each volume of Ecology foryears 1978-97 to assess the prevalence of trivialnull hypotheses and associated P-values in published ecological studies. We then randomlysampled 20 papers from each volume of theJournal of Wildlife Management (JWM) foryears 1994-98 for comparison. In each sampledarticle, we noted whether the null hypothesestested seemed at all plausible. In addition, wecounted the number of P-values and equivalentsymbols, such as statistics with superscripted asterisks or comparisons specifically marked nonsignificant. We tallied the number of caseswhere only a P-value was given (some papersalso provided the test statistic, degrees of freedom, or sample size), without an estimate ofeffect size, its sign or its precision, even in anassociated table, for papers appearing in theJWM during the 1994-98 period. However, ourcounts did not include comparisons that wereboth nonsignificant and unlabeled or unspecified, nor did they include all possible statisticalcomparisons or tests. Consequently, ours is anunderestimate of the total number of statisticaltests and associated P-values contained withineach article.In the 347 sampled articles in Ecology containing null hypothesis tests, we found few examples of null hypotheses that seemed biologically plausible. Perhaps 5 of 95 articles in JWMcontained -1 null hypothesis that could be considered a plausible alternative. Only 2 of 95 articles in JWM incorporated biological importance into the interpretations of results, the re-

916J. Wildl. Manage. 64(4):2000HYPOTHESISTESTING Anderson et al.Table 1. Median,mean (SE), and range of the numberof P-values per article,and estimatedtotal (SE) numberof P-valuesper year, based on a randomsample of 20 papers each year fromthe Articlessection of Ecologyfor 1978-97.Estimated no. of P-values per articleVolume yearTotal 52326x er merely used statistical significance. Inthe vast majority of cases, the null hypotheseswe found in both journals seemed to be obviously false on biological grounds even beforethese studies were undertaken. A major research failing seems to be the exploration of uninteresting or even trivial questions. Commonexamples included null hypotheses assumingsurvival probabilities were the same between juveniles and adults of a species, assuming no correlation or relationship existed between variables of interest, assuming density of a speciesremained the same across time, assuming netprimary production rates were constant acrosssites and years, and assuming growth rates didnot differ among individuals or species.We estimate that there have been a minimumof several thousand P-values appearing in everyvolume of Ecology (Table 1) and JWM (Table2) in recent years. Given the conservatism ofour counting procedure, the number of null hypothesis tests that were actually performed ineach study was probably much larger. Approximately 47% (SE 3.9%) of the P-values thatwe counted in JWM appeared alone, withoutestimated means, differences, effect sizes, or associated measures of precision. Such results, wemaintain, are particularly uninformative (e.g.,not even the sign of the difference being indicated). The key problem here is the general failure to explore more relevant questions and 1-2080-208Estimated yearly total (SE) 421)(945)(1,015)(1,720)(2,013)-- - - ----report informative summary statistics (e.g., estimates of effect size and their precision), evenwhen significance was found. The secondaryproblem is not recognizing the arbitrariness ofa, hence perpetuating an arbitrary classificationof results as significant or not significant.A PRACTICALALTERNATIVETO NULLHYPOTHESIS TESTINGWe advocate Chamberlin's (1890, 1965) concept of multiple working hypotheses rather thana single statistical null vs. an alternative-thisseems like superior science. However, this approach leads to the multiple testing problem instatistical hypothesis testing, and arbitrariness inthe choice of a-level and of which hypothesis toserve as the null. Although commonly used inpractice, significance testing is a poor approachto model selection and variable selection in regression analysis, discriminant function analysis,and similar procedures (Akaike 1974, McQuarrie and Tsai 1998:427-428).Akaike (1973, 1974) developed data analysisprocedures that are now called information theoretic because they are based on Kullback-Leibler (1951) information. Kullback-Leibler information is a fundamental quantity in the sciencesand has earlier roots back to Boltzmann's concept of entropy. The Kullback-Leibler information between conceptual truth, f, and ap-

HYPOTHESIS TESTING * Anderson et al.J. Wildl. Manage. 64(4):2000917Table 2. Median,mean (SE), and range of the numberof P-values per article,and estimatedtotal (SE) numberof P-valuesarticles)each year fromtheper year, based on a randomsample of 20 papers (excludingInvitedPapers and Comment/ReplyJournalof WildlifeManagementfor years 1994-98.Estimated number of P-values per 61041501662124212428. . . ,,,,,., , H ,Hm,.,x (SE)3237545631m,HH.proximating model g is defined for continuousfunctions as the integralI(f, g) f(x)loge(f(x)) dx,wheref and g are n-dimensional probability distributions. Kullback-Leibler information, denoted I(f, g), is the information lost when model gis used to approximate truth, f The right handside looks difficult to understand, however itcan be viewed as a statistical expectation of thenatural logarithm of the ratio off (full reality)to g (approximating model). That is, KullbackLeibler information could be written as/ f(x) lO))Ef logg(xwhere the expectation is taken with respect tofull reality, f Using the property of logarithms,this expression can be further simplified as thedifference between 2 expectations,I(f, g) Ef[loge(f(x))]- Ef[loge(g(x 0))].Clearly, full reality is unknown, but it is fixedacross models, thus a further simplification canbe written asI(f, g) C - EIloge(g(x10))],where the expectation of the logarithm of fullreality drops out into a simple scaling constant,C. Thus, the focus in model selection is on theterm Ey{loge(g(x O))].One seeks an approximating model (hypothesis) that loses as little information as possibleabout truth; this is equivalent to minimizing I(f,g), over the set of models of interest (we assumethere are R a priori models, each representingan hypothesis, in the candidate set). Obviously,Kullback-Leibler information, by itself, will notaid in data analysis as both truth (f) and theparameters (0) are unknown to us.(8)(10)(24)(16)(6),,,,,Estimated yearly total (SE) 6168,4005,146m,., j,m.H, ,m.,(808)(1,060)(2,496)(2,400)(996), H, H .Model Selection CriteriaAkaike (1973) found a formal relationship between Kullback-Leibler information (a dominantparadigm in information and coding theory) andmaximum likelihood (the dominant paradigm instatistics; deLeeuw 1992). This finding makes itpossible to combine estimation and model selection under a single theoretical framework-optimization. Akaike's breakthrough was derivingan estimator of the expected, relative KullbackLeibler information, based on the maximizedlog-likelihood function. This led to Akaike's information criterion (AIC),AIC -2log,(e(Ojdata)) 2K,where logfe(9 data) is the value of the maximizedlog-likelihood over the unknown parameters (0),given the data and the model, and K is the number of parameters estimated in that approximating model. There is a simple transformation ofthe estimated residual sum of squares (RSS) toobtain the value of log(fe(l(data)) when usingleast squares, rather than likelihood methods.The value of AIC for least squares models ismerely,AIC n.loge(&2) 2K,where n is sample size and &2 RSS/n. Suchquantities are easy to compute once the RSSvalues for each model are available using standard computer software.Assuming a set of a priori candidate models(hypotheses) has been defined and well supported, AIC is computed for each of the ap 1, 2,proximating models in the set (i.e., gi, i. . R). The model where AIC is minimized isselected as best for the empirical data at hand.This concept is simple, compelling, and is basedon deep theoretical foundations (i.e., KullbackLeibler information). The AIC is not a test inany sense: no single hypothesis (i.e., model) is

918J. Wildl. Manage.64(4):2000HYPOTHESISTESTING* Andersonet al.made to be the null, no arbitrary a level is set,and no notion of significance is needed. Instead,there is the concept of a best inference, giventhe data and the set of a priori models, andfurther developments provide a strength of evidence for each of the models in the set.It is important to use a modified criterion(called AICc) when K is large relative to samplesize n,2K(K 1)AICC -2 log(L (1 data)) 2K (n K 1)and this should be used unless n/K about 40(Burnham and Anderson 1998). As sample sizeincreases, AIC AICC,thus, if in doubt, alwaysuse AIC? as the final term is also trivial to compute. Both AIC and AICc are estimates of expected (relative) Kullback-Leibler informationand are useful in the analysis of real data in the"noisy" sciences.RankingModelsThe evidence for each of the alternative models can best be done by rescaling AIC valuessuch that the model with the minimum AIC (orAICc) has a value of 0, i.e.,Ai AICi - minAIC.The Ai values are easy to interpret and allow aquick strength of evidence comparison andscaled ranking of candidate models. The largerthe A., the less plausible is the fitted model i asbeing the best approximating model in the candidate set. It is generally important to knowwhich model (biological hypothesis) is rankedsecond best as well as some measure of itsstanding with respect to the best model. Suchranking and scaling can be done easily with theAi values.Likelihoodof a Model, Given the DataThe simple transformation exp(-hA), for i 1, 2, . ., R, provides the likelihood of themodel, given the data: 5(gi[data). These arefunctions in the same sense that S(Oldata, gi)is the likelihood of the parameters 0, given thedata (x) and the model (gi). It is convenient tonormalize these values such that they sum to 1,asexp(WiAi) Rra2- The wi, called Akaike weights, can be interpreted as approximate probabilities that modeli is, in fact, the Kullback

DAVID R. ANDERSON,'2 Colorado Cooperative Fish and Wildlife Research Unit, Room 201 Wagar Building, Colorado State University, Fort Collins, CO 80523, USA KENNETH P. BURNHAM,1 Colorado Cooperative Fish and Wildlife Research Unit, Room 201 Wagar Building, Colorado State

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