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A History of the Ecological Sciences, Part 9. Albertus Magnus: a Scholastic Naturalist
Medieval western Europe made a much greater invest-ment in higher education than any other civilization ever had, and science was a prominent part of the curriculum. There is no simple explanation as to why this happened, but a strong demand developed for scholars educated in theology, law, or medicine, and, of course this created a demand for professors to educate them. Italian universities, which were the earliest, tended to be sponsored by cities, although Frederick II founded the University of Naples in 1224 as a state university to educate state officials who were not clergy. Elsewhere, universities were usually church spon-sored, but even so, all universities in western Europe had much more autonomy than comparable institutions in other civilizations (Kibre and Siraisi 1978, Huff 1993: 149Ð201). A powerful stimulus to curriculum development was the translation of works by Aristotle and his Arabic commentators into Latin. Through much of the 1200s, there was a lively discussion between scholars and clergy over how appropriate it was to devote many courses to pagan learning. The Aristotelians, led by Albertus Magnus and his pupil Thomas Aquinas, pretty much won, although univer-sities did agree not to debate heretical questions, such as the possible eternity of the universe (Grant 1974:42Ð52). Albert the Great was from a noble family and was born about 1200 at the family castle of Lauingen. He grew up in the family manor in nearby Bollst‰dt, and was called Albert of Lauingen or Albert of Bollst‰dt. He studied lib-eral arts at the University of Padua, in Italy, and against the wishes of his family joined the Dominican Order. He was ordained in Germany and taught at several priories un-til he went to the University of Paris in 1240 or 1241. He earned an M.A. degree in theology and lectured there until 1248, when he went to Cologne to teach. Much of his time
from 1253 to 1262 was devoted to administrative duties for the Catholic Church, after which he returned to teaching, preaching, and writing until his death in 1280 (Wallace 1970). The Dominicans have a strong commitment to teach-ing, and his brothers in the Order asked him to explain, in Latin, AristotleÕs works. This task became his main lifeÕs work, and he was probably the most prolific author of the Middle Ages (Kitchell and Resnick 1999:18). He para-phrased all of Aristotle and added commentaries based upon his own observations and those of others. One of AlbertÕs earliest works isLiber de natura locorum, on geography. He reviewed the ancient arguments against the possibility of people being able to live at the Equator, but dismissed them because both Ptolemy and Ibn Sina had seen men who lived between the Tropic of Cancer and the Equator, and it was known that people live at the Equator in Africa and in the [East] Indies (Tilmann 1971: 54). However, he knew that life at 56 degrees latitude was difficult, and therefore he did believe that the poles were uninhabitable; they may have day for half a year and night for half a year (Tilmann 1971:65). Animals, such as bears and lions, that live in polar regions tend to be white. The sea freezes in winter and icebergs float in the sea in sum-mer (Tilmann 1971:67). Albert knew that the proximity of the sea modifies the climate of land, that high mountains can have perpetual snow, that mountains can also influence climate by blocking the wind, and that depressions of great depth can have noxious gas, as do swamps and some lakes (Tilmann 1971:86Ð89). He also thought that Òlands situated in the middle of great forests or near the forest, always have a suffocating and a thick air, and they have much fog and many whirlwinds.Ó It was not just forest in aggre-gate that was the problem, but also certain noxious trees: Òwalnut, the oak, and other trees which either by their bit-terness, poison the air, or by their height confine the air, and do not permit it to escape and be purified.Ó (Tilmann 1971:89Ð90). He also thought that living beings are influenced by their localities: mountains, seas, woods, swamps, and so on.
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ÒMen born in rocky places, level areas, and cold dry places are very strong and well-boned with visible joints. They are of noble stature, have skill and endurance in war, and have mus-cular limbs, and they have wild customsÉ.Ó On the other hand, those who live Òexposed to the south and not to other directions live poorly on account of the turbulent, warm, moist, and pestilential wind. Because their pores are opened on account of the heat, they must wear such clothing that will prevent the wind from penetrating to the marrow of their bones. Houses for them should be built with a strong protection from the southÓ (Tilmann 1971:106Ð107). The same was true for plants, animals, and stones. ÒThis is the proof. Bears in cold and moist places, and rabbits in places rather moist, cold, and dry [sic], are white, while in other climates they tend toward blackness, darkness, or they are golden yellowÓ (Tilmann 1971:108). He wroteDe vegetabilibus libri VII during the 1250s, based upon De plantisby Nicolaus of Damascus (first century BC), which he assumed was by Aristotle. He saw that it was not as well written or organized as AristotleÕs other works, but attributed this to a lack of understanding or skill by the translator (Alfred of Sarashel) from Arabic into Latin.De vegetabilibushas this organiza-tion: books I and IV paraphraseDe plantis; books II and V are AlbertÕs commentaries onDe plantis; book III summa-rizes a discussion by Ibn Sina on seeds, fruits, and fruit juices; book VI is a herbal describing some 400 species, in-cluding habitats and locations; and book VII is on agricul-ture, based largely onDe agriculturaRutilius Taurus by Palladius (late AD 300s). Book VII is Òthe best general work on agriculture since Columella [mid-first century AD] and shows how scientific thinking was stirred by the current technical changes in agricultureÓ (Morton 1981:93).  Albert was the only medieval encycloped-who added significant observations of his wn to supplement what he culled from his sources (Paszewski 1968, Stannard 1978,  1980a,b, Reeds 1980, Egerton 1983).  Book VI contains most of his personal  observations and is the book most ex- tensively translated into a modern lang- guage (Albertus Magnus 1992). He be-ieved that wheat and rye change from one ecies to the other, depending upon the soil in which the seeds are planted (Sprague 1933:432). His account of oak trees in-
Bulletin of the Ecological Society of America
cludes these observations and speculations on oak galls (Grant 1974:700):
On the leaves of the oak often grow certain round ball-like objects called galls, which after remaining some time on the tree produce within themselves a small worm bred by the corruption of the leaf. If the worm exactly reaches the midst of the gall apple, weather prophets foretell that the coming winter will be harder; but if it is near the edge of the gall, they foretell that the winter will be mild.
AlbertÕsDe vegetabilibusthe most important bo- was tanical work of the Middle Ages (Thorndike 1923:538Ð 539), yet it does not equal the botanical treatises by Theophrastos (c.371Ðc.287 BC), which he never saw. AlbertÕsDe animalibus libri XXVI was his most ex-tensive and influential work, probably begun between 1256 and 1260. It is organized into three parts: books IÐXIX paraphrase and explain the zoological works of Aristotle translated from Arabic by Michael Scot; books XXÐXXI are AlbertÕs additions to AristotleÑhis most original  contributionÑand books XXIIÐXXVI  are, translators Kitchell and Resnick  admit, a bestiary Òbased onDe  naturarerumof his former student,  Thomas of CantimprÈÓ (1999:40),  although this is not acknowledged  on the title page. AlbertÕs use of  ThomasÕs work deserves more than  passing mention. Thomas (1201Ð1270/  1272) was also of aristocratic birth, born at Lewes, near Brussels. He studied at LiËge, 1206Ð 1216, and then entered the Augustinian abbey at CantimprÈ. In 1232, Thomas transferred to the Domini-can Order and went to study under Albert at Cologne, 1233Ð1237; finally he studied atthe University of Paris, 1237Ð1240. While at Paris, he completed his encyclope-dia,De naturis rerumencyclopedia of all natu-Ñthe first ral phenomena in Latin since theNaturalis historia by Pliny (c. AD 23Ð79) (Ley 1968:92). Thomas was not a scholar like Albert, but rather a teacher who compiled popular scientific information for preachers to use for religious arguments in their sermons (Friedman 1974:107Ð 110, Kibre 1976). Nevertheless, Albert borrowed without acknowledgment most of this workÕs bestiary (Thomas of CantimprÈ 1973:101Ð311) for his own books XXIIÐXXVI. M. Bormans published this fact in 1852, and it was subse-quently accepted by several other scholars. The philologist Hermann Stadler (1906), however, contested this claim before publishing his edition of AlbertÕs Latin text (Albertus Magnus 1916Ð1921). There matters stood in 1931, when Pope Pius XI elevated Albert to the status of saint and doctor of the Church. Later, however, Pauline Aiken reexamined the matter and concluded that StadlerÕs article Òis a tissue of errorsÓ (1947:206). Aiken summa-rized her detailed findings (1947:225):
Albertus describes four hundred seventy-six specific creatures. For four hundred of these (more than five sixths of the total number) Thomas is the main source. In three hundred seventy-four of these descriptions (nearly four fifths of the total) there is either no supplementary material or not more than a few sentences per section.
Because Albert cited both ancient and Arabic sources, but not his contemporary Latin source, it is difficult to avoid the conclusion that he intentionally plagiarized from his former student. We are obliged to rectify the credits, but the influence of AlbertÕsDe animalibustaking justifies seriously this composite work. In their excellent translation ofDe animalibus, Kitchell and Resnick (1999) do not dis-tinguish, in books IÐXIX, between AlbertÕs paraphrase of Aristotle and AlbertÕs own additions (although this is often indicated in their notes), and their example is followed here. That distinction, however, is made in StadlerÕs edi-tion (Albertus Magnus 1916Ð1921). Next after his use of the works of Aristotle and Thomas, the third most impor-tant source for Albert seems to have beenDe animalibus by Ibn Sina (Avicenna, 980Ð1037), which, like AristotleÕs zoology, had been translated from Arabic into Latin by Michael Scot. The combined text and notes of Kitchell and ResnickÕs translation (Kitchell and Resnick 1999) are 1720 pages (StadlerÕs compact Latin text and notes [Stadler 1906] are 1598 pages)Ñtoo long for an adequate summary here of discussions having ecological interestÑyet it is pos-sible to indicate their character and scope (as Balss did in greater detail [1928:75Ð130], but not reprinted in Balss [1947]). The translatorsÕ introduction (Kitchell and Resnick 1999) emphasizes AlbertÕs skepticism of many fabulous reports in the scientific literature, his insistence upon natu-ral, rather than supernatural, explanations, and the many first-hand observations that he reported (Kitchell and Resnick 1999:36Ð42). This is all true, and is why hisDe animalibusof lasting interest. However, it is also true is that there were limits to his ability to discriminate, and there were reports, which he accepted, that are no longer credible. Books IÐIV are on the anatomy and physiology of animals and humans, topics only indirectly relevant to eco-logical sciences, but it is interesting methodologically to follow his attempt to determine whether Aristotle is cor-rect in saying that veins and arteries both arise at the heart, or Galen is correct in saying that veins arise at the liver and arteries at the heart. The answer presumably had im-portant implications for physiology. To resolve the matter, Albert not only discussed the arguments of both men, he also brought in those of Ibn Sina and Ibn Rushd (Averroes, 1126Ð1198). Albert believed that he then could reach a conclusion Òby use of reason and solid experiential knowl-edge that is completely trustworthyÓ (Tilmann 1971:351). He concluded that Aristotle was correct, but he did so by using Aristotelian logic. If he based any of his arguments upon his own observations, he failed to say so, and Òthe
solid experiential knowledgeÓ that he referred to appar-ently was obtained by Ibn Sina and Ibn Rushd. Books VÐVI are on animal reproduction and books IXÐ X are on human reproduction. He accepted the Aristotelian idea that some animals are created from mud, putrescence (Tilmann 1971:516), slime, or sand, and Òthey differ to the extent that the slime or sand from which they are generated differsÓ (Tilmann 1971:518Ð519). After a survey of the re-production of oceanic nonfish animals [invertebrates], however, he acknowledges that ÒIt is not possible to know the differences in generation of all these animals well, be-cause the experts can scarcely observe the times of the conception, egg laying, and emergence of the young of these animalsÓ (Tilmann 1971:524). He mentions the fact that different species of birds lay different numbers of eggs without much speculation as to why. For eagles, however, he reported that they lay three eggs but only raise two chicks, and does offer an explanation: ÒSome say the rea-son for this is that it is so weakened by incubating the eggs that it cannot hunt enough chicks of other birds for three and is scarcely able to care for two of themÓ (Tilmann 1971:545). In book VIII, ÒOn AnimalsÕ Habits,Ó Albert added this to AristotleÕs discussion of hawks and eagles (Tilmann 1971:599):
Of all the genuses of eagle and falcon, the best and the fiercest is the one which comes from the northern region of Sweden and Latvia whose latitude is more than fifty degrees from the equator. These are fierce birds and they would rather eat fish than flesh. Thus, certain astures which were brought from that land to our land all catch birds to be sure, but they eat crabs more readily than any other food. These astures are held by the falconers in our land to be better and nobler than any others, and they are very large. One who is quite an expert said to me that even in that land the eagles mostly feed on fish and that eels and fish are thus found in and near their nests.
AlbertÕs discussions of hawks and falconry have been closely studied, including his sources. We know that he borrowed from Thomas of CantimprÈ, but did he also have access to Frederick IIÕsDe arte venandi cum avibus? Ap-parently not: Òit seems more likely that Albert was trans-mitting information passed on to him orally by FrederickÕs
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falconersÓ (Oggins 1980:444). Regardless of his sources, AlbertÕs discussion seems generally well informed. How-ever, it also includes this folklore: ÒThere is in our land, also a small genus of eagle called the fish-eagle [osprey]. It hunts only fish and has one webbed foot, for swimming, like that of a goose, while the other foot has hooked talons for seizing, like that of an eagleÓ (Tilmann 1971:599). Albert was widely traveled in Europe (maps: Kitchell and Resnick 1999:7, 9, Tilmann 1971:19), and this experi-ence may have both increased his interest in migrations and also added to his knowledge of it. He knew that heat or cold that was healthful to some species could be harmful to others. He thought that migration was due either to heat (in spring) or cold (in autumn), although he knew of vari-ous ways in which animals responded to changes in cli-mate (Tilmann 1971:613):
Of those animals which do go away, some go to elevated places, seeking in them a temperate cold. Others, however, go to cave-like, warm places, seeking warmth in them. They hide themselves away in the same placesÑin hollows in trees or beneath drooping leaves or in actual cavesÑseeking heat. Some do not change place. In our lands the cranes are present continuously in winter and summer. Still, our dwelling place is very cold, being almost 47 degrees of latitude from the equator.
Geese bred in Sclavia at Òmoist, sandy, marshlike places,Ó but at the beginning of winter they Òcome back to our land,Ó which he said was at a latitude of 47∫, seeking Òfood and the more temperate airÓ (Tilmann 1971:613). They fly south on winds from the north, in flocks of thou-sands. Fish either migrate or seek out holes for the winter. ÒSome fish move from the depths of the sea during the winter and come near the landÕs edge in search of heat. Others do the opposite and flee the shore for the depth of the sea, escaping the shoreÕs heatÓ (Tilmann 1971:614). Albert also claimed that when animals migrate from a warm to a cold place, they fatten, but when they return they grow thin Òbecause of the dissolving and consuming heatÓ (Tilmann 1971:615). Discussions of differences between the sexes were susceptible to gender bias and/or folklore, and Albert suc-cumbs to some of those reports. He reported that, in gen-
Bulletin of the Ecological Society of America
eral, females are easier to train than males, and this is espe-cially true among dogs (Tilmann 1971:668). Among quad-rupeds, females are weaker than males, but in birds of prey females are the larger and stronger sex. A female bear Òhas boldness due to the bad habits which are attendant upon her sexÓ (Tilmann 1971:669). Females of almost all spe-cies Òare fiercer than the males during the time in which they have young.Ó As for humans, he cited Ibn Sina to the effect that women are Òstupider when it comes to honest and good things, and to governanceÓ (Tilmann 1971:669). Presumably, he wrote only for male readers, because it did not seem necessary to support this claim with evi-dence. Moving on to mating, he reported that pigeons re-main faithful to each other after mating, but that Ibn Sina saw two males fight over a female, which accepted the winner, but when the loser returned to fight again and this time won, the female accepted him instead. After copula-tion, Òthe female follows the male and obeys him. When, however, the female does not come into the nest quickly the male beats her with his wingsÓ (Tilmann 1971:689). However, even on sexual matters, there were limits to what Albert could believe. He could believe that a female turtledove remained faithful to her mate during his life-time, but not afterwards: ÒSome say that even after the death of this one she does not take another husband, but this is neither probable nor has it been verified through ex-perienceÓ (Tilmann 1971:690). Albert was the outstanding encyclopedist of the High Middle Ages (1000Ð1350);De natura locorum was first printed in 1514,De vegetabilibus1517, and in De animali-busin 1478. There were other encyclopedists whose works were as widely or more widely read than his. At best, all of their encyclopedias blended fact and folklore, and AlbertÕs had the most first-hand information. There was no simple progression toward greater and greater accuracy from one to the next. Like Islamic civilization, medieval Europe suffered a catastrophe and decline; the Black Death struck in 1347 and continued, off and on, for several centuries. It is generally understood to have been bubonic plague, although recent scholarship suggests that the original epi-demic might not have been limited to just one type of in-fection (Cantor 2001:part I). Unlike Islamic civilization, how-ever, Europe rebounded a century later, stronger than ever. The survival of universities and GutenbergÕs invention of the printing press were important factors in that recovery.
I thank Anne-Marie Drouin-Hans, UniversitÈ de Bourgogne, and Jean-Marc Drouin, MusÈe Nationale dÕHistoire Naturelle, for their suggestions.
Literature cited
Aiken, P. 1947. The animal history of Albertus Magnus and Thomas of ContimprÈ. Speculum22:205Ð225. Albertus Magnus. 1867. De vegetabilibus libri VII, historiae naturalis pars XVIII. E. H. F. Meyer and K. Jessen, edi-tors. G. Reimeri, Berlin, Germany. Albertus Magnus. 1916Ð1921. De animalibus libri XXVI. H. Stadler, editor. Aschendorffsche Verlagsbuchhand-lung, M¸nster, Germany. Albertus Magnus. 1992. De vegetabilibus Buch VI, Traktat 2. Latin with German translation by K. Biewer. Wis-senschaftliche Verlagsgesellschaft, Stuttgart, Germany. Albertus Magnus. 1999. On animals: A medieval summa zoologica. Translated and edited by K. Kitchell, Jr., and I. M. Resnick. Two volumes. Johns Hopkins University Press, Baltimore, Maryland, USA. Balss, H. 1928. Albertus Magnus als Zoologe. M¸nchner Drucke, M¸nchen, Germany. Balss, H. 1947. Albertus Magnus als Biologe: Werk und Ur-sprung. Wissenschaftliche Verlagsgesellschaft, Stuttgart, Germany. Cantor, N. F. 2001. In the wake of the plague: the black death and the world it made. Free Press, New York, New York, USA. Egerton, F. N. 1983. Latin Europe. Pages 444Ð452inF. N. Egerton, editor. Landmarks of botanical history. Two volumes. Stanford University Press, Stanford, Califor-nia, USA. Friedman, J. B. 1974. Thomas of CantimprÈ. De naturis rerum prologue, Book III and Book XIX. Cahiers DÕ…tudesMÈdiÈvales2 (La science de la nature, thÈories et pratiques):107Ð154. Grant, E., editor. 1974. A source book in medieval science. Harvard University Press, Cambridge, Massachusetts, USA. Huff, T. T. 1993. The rise of early modern science: Islam, China, and the West. Cambridge University Press, New York, New York, USA. Kibre, P. 1976. Thomas of CantimprÈ, also known as Tho-mas Brabantinus. Dictionary of Scientific Biography 13:347Ð349. Kibre, P., and N. G. Siraisi. 1978. The institutional setting: universities. Pages 120Ð144inC. Lindberg, editor. D. Science in the Middle Ages. University of Chicago Press, Chicago, Illinois, USA. Kitchell, K. F., and I. M. Resnick. 1999. Introduction: his life and works. Pages 1Ð44inAlbertus Magnus, on ani-
mals: A Medieval Summa Zoologica. Johns Hopkins University Press, Baltimore, Maryland, USA. Ley, W. 1968. Dawn of zoology. Prentice-Hall, Engle-wood Cliffs, New Jersey, USA. Morton, A. G. 1981. History of botanical science: an ac-count of the development of botany from ancient times to the present day. Academic Press, London, UK. Oggins, R. S. 1980. Albertus Magnus on falcons and hawks. Pages 441Ð462in J. A. Weisheipl, editor. Albertus Magnus and the sciences: commemorative essays. Pontifical Insti-tute of Medieval Studies, Toronto, Ontario, Canada. Paszewski, A. 1968. Les problËms physiologiques dans De vegetabilibus et plantis libri VII dÕAlbert von Lauingen. Actes du XIe CongrËs International dÕHistoire des Sciences5:323Ð330. Reeds, K. 1980. Albert on the natural philosophy of plant life. Pages 341Ð354inJ. A. Weisheipl, editor. Albertus Magnus and the sciences: commemorative essays. Pon-tifical Institute of Medieval Studies, Toronto, Ontario, Canada. Sprague, T. A. 1933. Plant morphology in Albertus Magnus. Kew Bulletin of Miscellaneous Information1:440Ð 459. Stadler, H. 1906. Albertus Magnus, Thomas von CantimprÈ, und Vinzenz von Beauvais. Natur und Culture4. Stannard, J. 1978. Natural history. Pages 429Ð460inD. C. Lindberg, editor. Science in the Middle Ages. Univer-sity of Chicago Press, Chicago, Illinois, USA. Stannard, J. 1980a. Albertus Magnus and medieval herbal-ism. Pages 355Ð377inJ. A. Weisheipl, editor. Albertus Magnus and the sciences: commemorative essays. Pon-tifical Institute of Medieval Studies, Toronto, Ontario, Canada. Stannard, J. 1980b. The botany of St. Albert the Great. Pages 354Ð373inG. Meyer and A. Zimmermann, edi-tors. Albertus Magnus: Doctor Universalis, 1280/1980. Grˆnewald, Mainz, Germany. Thomas of CantimprÈ. 1973. Liber de natura rerum. H. Boese, editor. W. De Gruyter, Berlin, Germany. Thorndike, L. 1923. Albertus Magnus. Volume 2:517Ð592. A history of magic and experimental science. Columbia University Press, New York, New York, USA. Tilmann, J. P. 1971. An appraisal of the geographical works of Albertus Magnus. University of Michigan Press, Ann Arbor, Michigan, USA. Wallace, W. A. 1970. Saint Albertus Magnus. Dictionary of Scientific Biography1:99Ð103.
Frank N. Egerton Department of History University of Wisconsin-Parkside Kenosha WI 53141 E-mail: frank.egerton@uwp.edu
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Normality of Raw Data in General Linear Models: the Most Widespread Myth in Statistics
In years of statistical consulting for ecologists and wildlife biologists, by far the most common misconception we have come across has been the one about normality in general linear models. These comprise a very large part of the statistical models used in ecology and includettests, simple and multiple linear regression, polynomial regres-sion, and analysis of variance (ANOVA) and covariance (ANCOVA). There is a widely held belief that the normal-ity assumption pertains to the raw data rather than to the model residuals. We suspect that this error may also occur in countless published studies, whenever the normality as-sumption is testedprioranalysis. This may lead to the to use of nonparametric alternatives (if there are any), when parametric tests would indeed be appropriate, or to use of transformations of raw data, which may introduce hidden assumptions such as multiplicative effects on the natural scale in the case of log-transformed data. Our aim here is to dispel this myth. We very briefly de-scribe relevant theory for two cases of general linear mod-els to show that the residuals need to be normally distrib-uted if tests requiring normality are to be used, such ast andFtests. We then give two examples demonstrating that the distribution of the response variable may be nonnormal, and yet the residuals are well behaved. We do not go into the issue of how to test normality; instead we display the dis-tributions of response variables and residuals graphically.
A very brief theory of general linear models
We present two simple examples from among the large class of general linear models, which encompass such meth-ods as, e.g., thet test, simple and multiple linear regres-sion, polynomial regression, ANOVA, and ANCOVA. In every case, a response variable is thought to be composed of additive systematic components and one or several ran-dom components. The latter are usually assumed to be from a common normal distribution with a constant variance.
Simple linear regression
The normal error regression model for a sample of size nthat links a response variateYto one continuous explana-tory variableXis as follows (from Neter et al. 1990:52):
Y=β+βX+ε i0 1i i
whereYis the observed response for theith unit;Xis the i i value of the explanatory variable for theith unit;βandβ 0 1 are parameters, i.e., unknown constants to be estimated 2 from the data;εindependent N(0, are σ) , i.e., indepen-i dent normally distributed residuals about a zero mean with 2 constant varianceσ;i= 1,É,nand indexes the units.
Bulletin of the Ecological Society of America
Normality and homoscedasticity (constant variance) of residuals is not necessary to use the least-squares or maxi-mum-likelihood method to provide unbiased point esti-mates of the parametersβ andβ. However, to provide 0 1 significance tests or confidence intervals, the standard as-sumption of a normal distribution of error termsεneeds to i be invoked (Neter et al. 1990).
One-way ANOVA
The linear additive model links a response variateYto one discrete explanatory variable withI(discrete levels values) and can be written as (from Steel and Torrie 1980:149):
Y=µ+τ+ e ij i ij
whereYis the observed response for thejth unit in group ij i;µandτare unknown constants, i.e., parameters to be es-i timated from the data;µis the overall mean response andτ i is the additive effect of leveli(i= 1,É,Iare indepen-); e ij dent random components;j= 1,É,nand indexes the units within each level of the explanatory variable. Again, when significance tests or confidence intervals are desired, distributional assumptions about the random components e need to be made. Customarily, they are as-ij sumed to be independent normally distributed with zero 2 mean and constant varianceσ.
Two numerical examples
Multiple linear regression
The number of fruits per stem ofGentiana cruciata, a rare plant of calcareous grasslands, had been measured on 810 plants and ranged from 1 to 60. The distribution of these data was clearly not normal, but right skewed (Fig. 1a). A multiple-regression model using three continuous explanatory variables (population area, number of stems per plant, and length of the longest stem) fit by the package Genstat (Payne et al. 1993) accounted for 47% of the vari-ance in the data. It showed that the response variable was significantly and positively related to all three explanatory variables. The residuals of this model were reasonably close to a normal distribution (Fig. 1b). These data are from a larger study on a rare plant and its specialist herbi-vore. (For further description of the system, see KÈry et al. 2001.)
One-way ANOVA
We then generated 200 data points for each of four populations. Think of it as the mean number of seeds per fruit ofGentiana cruciata. Mean numbers of seeds were 100, 200, 300, and 400 in the four populations, respec-tively. Normally distributed noise with variance 50 was added. The distribution of these data was again far from normal (Fig. 1c). However, when the systematic popula-
Fig. 1.Distribution of the raw data and of the residuals (a,b) for a multiple linear regression analysis, and (c,d) for a one-way analysis of variance.
tion effect was taken out by fitting one discrete explana-tory variable, the resulting residuals were normally dis-tributed (Fig. 1d), as would be expected in this simu-lated case.
There is a very widespread misconception that, in gen-eral linear models, the raw data instead of the residuals of a model have to be normally distributed to permit con-struction of confidence intervals and significance statistics. Here we state this to be false and give two examples that show raw data may have some other distribution, and yet the residuals of a linear model turn out to be reasonably close to a normal distribution. Such examples abound, and we think that only in a minority of cases are the raw data already clustered symmetrically around a single mode. Re-sidual analysis for linear models is easily conducted in the two statistical packages that we are familiar with, Genstat (Payne et al. 1993) and SAS (SAS 2001). Residuals are easily stored in each analysis and then examined visually, e.g., by histograms or plots, or by formal statistical tests for normality. There is a large literature on model check-ing, also called model criticism, in general linear models (e.g., Cook and Weisberg [1982], Atkinson [1985], and also general texts on regression such as Draper and Smith
[1981], or Neter et al. [1990]). Model criticism is an im-portant part of any statistical modeling. In summary, we hope that this note is a contribution toward better statis-tical practice by doing away with the myth of normality of the raw data in general linear models.
Literature cited
Atkinson, A. C. 1985. Plots, transformations and regres-sion. An introduction to graphical methods of diagnos-tic regression analysis. Oxford University Press, Ox-ford, UK. Cook, R. D., and S. Weisberg. 1982. Residuals and influ-ence in regression. Chapman and Hall, London, UK. Draper, N. R., and H. Smith. 1981. Applied regression analysis. Wiley, New York, New York, USA. KÈry, M., D. Matthies, and M. Fischer. 2001. The effect of plant population size on the interactions between the rareGentiana cruciata and its specialized herbivore Maculinea rebeli. Journal of Ecology89:418Ð427. Neter, J., W. Wasserman, and M. H. Kutner. 1990. Applied linear statistical models. Third edition. Irwin, Burr Ridge, Illinois, USA. Payne, R. W., P. W. Lane, P. G. N. Digby, S. A. Harding, P. K. Leech, G. W. Morgan, A. D. Todd, R. Thompson, G. Tunnicliffe Wilson, S. J. Welham, and R. P. White.
April 2003
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Marc KÈry and Jeff S. Hatfield USGS Patuxent Wildlife Research Center 11510 American Holly Drive Laurel, MD 20708 (301) 497-5632 Fax: (301) 497-5666, E-mail: mkery@usgs.gov
What is Conservation Ecology? Conservation Ecologyis an electronic, peer-reviewed, multidisciplinary journal devoted to rapid dissemination of current research. Manuscript submission, peer review, and publication are handled on the Internet. Software developed for the journal automates all clerical steps, facilitates a double-blind peer review process, and allows authors and editors to follow the progress of review on the Internet. As articles are accepted, they are published in an “Issue in Progress.” At six-month intervals, the Issue-in-Progress is declared a New Issue. Subscribers receive the Table of Contents of the issue via e-mail.
What types of articles do we publish? We seek papers that are novel, integrative, and accessible to a wide audience from an array of disciplines (biology, ecology, economics, and social sciences) and concerned with issues of conservation, sustainability, development, and ecological policy. Content ranges from applied to theoretical. In general, papers should cover topics related to ecological, political, and social foundations for sustainable socioecological systems including: conservation, management, and sustainable use of ecological systems, resources, and biological diversity; the role natural systems play in social and political systems and vice versa; and the means by which we can develop and sustain desired ecological, social, and political states. We encourage papers using the unique opportunities of an e-journal: color illustrations, animated model output, downloadable models and data sets, a “Response” option for interactive discussion, and other inventions encouraging reader interaction. Our audience Conservation Ecology’saudience is an international community of scholars and practitioners representing the natural and social sciences, business, government, and NGOs. Most of the 11,000+ subscribers are affiliated with academic institutions, 31% represent commercial organizations, and 4% nonprofit organizations. Of the ~3250 U.S. subscribers, 17% are affiliated with government organizations. With subscribers in 108 countries (>50 developing countries) and many more regular “un-subscribed” readers, Conservation Ecologybridges disciplines and facilitates communication among researchers and policy makers, providing a mechanism for sharing more international perspectives. Nearly two-thirds of our subscribers hail from North America (USA 45%, Canada 11%); the remaining third is well distributed (Western Europe 13%, Latin America 11%, Australasia 8%, Scandinavia 3%, Central/Northern Asia 2%, Conservation Ecologyis included in Southeast Asia 2%, Southwest Asia1%, Africa 2%, Eastern Europe 2%, the the Expanded Science Citation Index Caribbean 0.5%). Easy and rapid international access to the journal is provided (Institute for Scientific Information), by mirror sites in Sweden and South Africa. The web site averages 140,000 hits in Current Contents (Agriculture, Bio-per month. Web Server Statistics indicate that we serve requests from >70,000 logy, and Environmental Sciences), distinct hosts, suggesting a much broader audience than is revealed by our list and Alerting Services. Abstracts are of subscribers. Subscription is not required but is free. It includes: e-mail notice in BIOSIS (Biological Abstracts and of each new issue, special papers and features, news from Resilience Alliance Zoological Records), Cambridge Sci-Projects, and notices of research writing competitions. entific Abstracts (Biological Sciences database), and Public Affairs Informa-Publisher:Resilience Alliance, Carleton University, Ottawa, Canada tion Service (PAIS International). Founding Editor-in-Chief:C. S. Holling Editors-in-Chief:Carl Folke and Lance Gunderson E-mail:questions@consecol.org •Phone:+ 1 613 520-3657http://www.consecol.org
Bulletin of the Ecological Society of America