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P RE!" T ICE- IIM .L J!'iT ER N ATIO NAL , I:-Ie ., Lon don PRENTICE-HALl.. OF AUSTRA LIA. P T Y. , LTD " Sydney PR EN TICE- HALL OF C'Ar\AOA, LTD., Toronto PR ENTICE-HALL OF l :-< lJI A (P RIVATE) LTD., Ne w De/hi PR E:--ITl CE· Il ALT. 01' JAI' AJ' , INC., Tok yo Current printing (la st digit) : 11 10 9 e 1965 by PRE:" n CE -HALL, l N C. , Engle wo od Clif fs, N ew Jersey A 1/ righ ts reserved. N o pari of this boo k may be repr oduced ill any form , by mim eograph or all y Ot/I ('/' menus, without permission in writing from the publ isher. Library of COII grl' SS Cutalog Card N umber: 65- 26924. Print ed in the United States of America. 75586-C PREFACE Thi s group of papers represents as wide a range Of the ecological literature as could be encompassed within a reasonably sized volume . Although' it is compiled as a textbook supplement for the beginning student in ecology, certain papers are suitable for college students in beginning biology or general science and for high school students in advanced biology . For the graduate student and research ecologist it provides an opportunity fa become reacquainted with the sources of their endeavors. With few exce ptions, all of these papers have been used in my course in general ecology at Oberlin College because of the trem en- dous potential inherent in ex posure to the original literature. Ai- though I have used them largely in synchrony with textbook declamations, their real value lies in the excitement and flavor, and the relating of the adve nture, hard work , and creativity which is science . Th ere is amp le testimony that these savors are expe rie nced and that scientific literatur e is co mprehensible. Th e criteria which guided the filtering out 'of few pap ers from the myriad for my own students have operated in meeting the exigencies im posed by publica/ion. Th e prime criterion is a pap er whi ch lias made a highly significant contribution; this is fairly ,9 10 5

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Page 1: Lon don PREFACE - people.uncw.edu

P RE!" T ICE- IIM .L J!'iT ER NATIO NAL, I:-Ie ., Lon don P RENTI CE- HALl.. OF AUSTRA LIA. PT Y. , LTD " Sydney

PR EN TICE- HALL OF C'Ar\AOA, LTD., Toronto PR ENTICE-HALL OF l:-< lJI A (P RIVATE) LTD., Ne w De/hi

PR E:--ITlCE· Il ALT. 01' JAI'AJ' , I NC., Tok yo

Current printing (la st digit) :

11 10 9

e 1965 by PRE:"n CE -HALL, l N C., Englewood Cliffs, N ew Jersey

A 1/ righ ts reserved. N o pari of this boo k may be reproduced ill any form , by mimeograph or ally Ot/I('/' menus, without perm ission in writ ing fro m the publisher. Library o f COII grl'SS Cutalog Card N umber: 65- 26924. Print ed in the United States of America. 75586-C

PREFACE

Thi s group of papers represents as wide a range Of the ecological literature as could be encom passed within a reasonably sized volume . A lthough' it is compiled as a textbook supplement for the beginning student in ecology , certain papers are suitable for college students in beginning biology or general science and for high schoo l students in advanced biology . For the graduate student and research ecologist it provides an op portunity fa become reacquainted with the sources o f their endeavors.

With few exce ptions, all o f these papers have been used in my course in general ecology at Oberlin College because of the trem en­dous potential inherent in ex posure to the original literature. Ai­though I have used them largely in synchrony with textbook declamations, their real value lies in the excitement and flavor, and the relating of the adve nture, hard work , and creativity which is science . Th ere is ample testimony that these savo rs are experienced and that scientific literatur e is comprehensible.

Th e criteria which guided the filtering out 'of few pap ers from the myriad for my own students have operated in meeting the exigencies im posed by publ ica/ion. Th e prime criterion is a pap er which lias made a highly significant contribution; this is fairly

,9 10 5

borretts
Typewritten Text
Kormondy, E.J. 1965. Readings in Ecology. Prentice-Hall, Inc. NJ. 219 pp.
Page 2: Lon don PREFACE - people.uncw.edu

readily attested by the frequency of its citation in subsequent liter­ature. In m ost instances, this standard also placed in the list m ost of the ecologists who ha ve given direction to their field . Other papers were selected as exem plary of good scientific procedure (trenchant analysis, incisive logical argument , stim ulating theoretical discourse) and others to achieve an indication of the range of eco­logical investigation,

Each paper has been abridged (indicated by an ellipse, ' . .) and had literature citations om itted primarily to conserve space. Th ere are recognized dangers here, not the least is that the reader may not be made fully aware of the dependence of a scientist on his predecessors and contem poraries.

Noble criteria notwithstanding, there is the hazard of autocracy in compiling a set of papers, perhaps more so for so div ersified a field as ecology. No two ecologists would likely develop identical lists nor present them in the order used here. M ost would concur in certain papers ha ving classic status, but at that juncture diver­gence of considerable magnitu de would begin. Th ere is an historic and functional reasonableness to both the selections and order used but not such as to deny great flexibility in selecting papers to meet individual requirem ents.

Because one's perspectives are always circumsc ribed and his hindsight want ing , the ut llity o f this volume call be increased in the future through criticism . charitable and otherwise , by its users. 1 will appreciate receiving tha i criticism .

Each of my teachers, students and colleagues (in person and, through the literature) has contributed in some way to the develop ­ment of this volum e. None is singled out since each is completely absolved from the final errors of com mission and omission, all of which arc m ille. I do, however, acknow ledge the coo peration of the authors and publishers wh ose permission made the volume possible, the translations by Mrs. Ursula Stechow and Dr. Lawrence Wilson, and the invaluable and diversified help of the library staj] and facilities at Oberlin College, the South Branch of the Univer­sity of Georgia, and the Savannah Ri ver Laboratory, 1 am con­tinually grateful to my wife, Peggy, for her editorial assistance but more for her encouragem ent and understanding. A sabbatical leave from Oberlin ( t 963-64) provided the time to com plete this under­tak ing,

EDWARD J . KORMON DY

CONTENTS

EARLY NATURAL HISTORY

TH EOPH RASTUS-C. 30 0 B. C.

E nquiry into plants 2

Ll !':N AE US-/ 7 5 0

History of plants 4

RE:--r t AN TO IN E F ERCH AULT DE REAUMUR-

Th e nat ural history of ants C, / 7 4 2

8

m E PHYSICAL AND CHEMICAL ENVIRONMENT

A nalysis of Environme ntal Factors

.J L'STUS LIEBlG-J 840 Organic chemi stry in its appli cation to vegetable physiology and agricultu re 12

F . F . RL ACK:\1AN- J 9 0 5

Optima and limiting factors vi

vii

14

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VICTOR E . SHELFORO- 191 1

Physiological anim al geograp hy 17

LAWRENCE J. HENDERSON-1913

Wat er-general considerations; thermal properties 20

RU DOL F GEIGF R- 194 1

H eat exc hange near the ground; cold air floods and cold air dam s 24

GEORGE L. CI.ARKE-1939

Th e utili zation of solar energy by aquatic organisms 27

Regulation of Organisms by Environment

HENR Y J . OOSTING AND W. DW IGH T BII.UNGS-1942

Factors effecting vegetational zonation on coastal dunes 3 1

W. W . GARN ER AND H. A. ALLARO- 1 9 2 3

Further studies in photop eriodism, the response of the plant to relative length o f day and night 3 6

Compensatory and Behavioral Adjustments to En vironment

T HEO DORE H . BU L L OCK-1 9 5 5

Compensation for temp erature in the metabolism and acti vity of poikil otherms 40

HERB ERT S. J ENNI NGs--1904

Th e method of trial and error in the behavio r of lower organisms 4 4

JACQUES LO EB- 1918

Forced movements, tropism s and animal conduct 47

DOUG L AS A. SPALDING-1872

Instinct , witn original observa tions on young animal s 49

KONRAD L OREN Z AN D NIKO TINllERGEN- 1 938

Taxis und instinkthandlung in del' eirollbewegung del' Graugans. 1. 53

ARTHUR D. H ASL ER AN D WARR EN J . W ISBy-1951

Discrimination of stream odors by fishes and its relation to parent stream behavior 56

viii

THE STUDY OF POPULAnONS Nature of Po pulatio n Growth

THOMAS ROBER T M AL THlJS-1 7 9 8

A n essay on the principle of po pulation as it affec ts the future im prove ment of society 6 2

P . F . VERHULS T ­ 18 3 8

Notice sur la loi qu e la pop ulation suit dans son accroisem eru 64

RAY M OND PEARL AN D LO WELL J. REF.D-1 9 2 0

On the rate of growt h of the population of the United Stat es since 1790 and its math emat ical representation 66

ROYA L N. CH APM AN-1 92 8

The quantitative analysis of env ironment al factors 69

CHAR L E S S. ELTON­ 1924

Periodi c fluctuations in the numb ers of animals: th eir causes and effec ts 73

Int eraction of Po pulations

PAUL R. BURKHOLDER-1952

Cooperation and conflict among prim iti ve organisms 77

G. F. GAUSE- I934 Co m petition for com mo n food in protozoa 82

.JOSEPI] H. CON NELL­ 196 l

Th e influence of intersp ecific competition and other factors on the distribut ion of the barnacle Chtharnalu s stella tus 86

H. E I.IOT lIOW ARD--1 9 2 0

Territory in bird life 90

WI LLIAM MORTON W H EELER-1 9I8

A study of some ant larvae with a consideration of the origin and m eaning of the social habit am ong insects 9 4

w. c . A I. LEE­ 1926

Studies in. animal aggregations: causes and effects of bun ching in land isop ods 98

R egulation of Pop ulations

DAVID LA CK-1948

N atural selection and fami ly size in the starling

ix

102

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L. C. BIRCH-1957

The role ot weath er in determining the distribution and abundance oj anim als 106

A. J. NICH OLSON -1 9 57

Th e seli-adiust me nt of populations to change 109

NELSON G. HAIRSTON, FREDERICK E. SMITH, AND

L AWREN CE B . SLOBODKTN - 1 9 6 0

Community structure , population con trol, and com petition 113

TIlE STUDY OF COMMUNITIES

Characteris tics of Communities

F.DW ARD FORB ES-I844

Report on th e M ollu sca and Radiata on the IF.gean Sea, and on the ir distribut ion co ns ide red as hearing on geology 118

KARL MOBIUS-I 877 A n oyster-bank is a biocon ose, or a social community 121

J,UGEN E Wi\RMING-IY0 9

Oecology oj plants, an introduction to the study oj plant com munities 1 25

H ENRY C. COWLES-18 99

The ecological relations o f the vegetation on the sand dun es of Lake Mi chigan 129

CAL VIN MCMJI.LAN-1959

The role of ecoty pic variation in th e distribution oj the cen tral grassland of N orth A m erica 133

EDW ARD S. DEEV EY, J R.- 1939

Studies on Connecticut lak e sediments . 1. A pas/glacial climatic chrono logy for southe rn N ew England 137

Nature oj th e Community

FR EDERIC E . CL EMENTS-1916

Plant succession, an analysis oj the development o f vegetation 140

V. N . SlJKATCHE W- /928

Prin cipl es oj classification of the spruce communities of European Russia 144

.JOSIAS IlR A UN -IlLAN()UET AN D ER:--: ST F CRRER-191 3

Remarques sur l'etude des groupements de plantes 147

x

L. G. RAMENSKY-1926 Die Grundgesetzmiissigkeiten im A ujbau der V egetationsdecke 151

H. A. GLE ASON-192 6 The individualistic concept oj th e plant association 153

STANLEY A. CAIN-1 947 Characteristics o f natural areas and factors in th eir development 157

R. H . WHlTTAKER-1953

A consid eration oi climax theory : th e clim ax as a population and pal/ern 159

THE CONCEPT OF THE ECOSYSTEM Th e Ecosystem Concept

FRANCIS C. EVANS-1956

Ecosystem as th e basic unit in ecology 166

ST EPHEN A . FORB ES-18 87

The lake (2'; a microcosm 168

Energetics oj E cosyst ems

EDGAR N ELSON T RANSEAU - 1926

The accumulation oj energy by plants 171

CHAN CEY JUDAY-1940

Th e annual energy budget o j an inland lak e 174

RAYMOND L. LlNDEM AN-1942

The trophic-dynamic asp ect o f eco logy 179

GEORGE L . CLARKE­ 19 46

Dynamics oj production in a marin e area 1 84

HOWARD T . ODUM-1957 Tr ophic structure and produc tivi ty of Sil ver Springs, Florida 188

B iogeoch em ical Cy cling

II. W. HARVEY-1926

Nitrate in the sea 191

xi

O . A RR IJENlUS­ 1922

Hydrogen ion conc entratio n, soil prop ert ies and growth oj higher plants 193

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ALFRED C. RELJFIELD--I958

Th e biological control of chem ical factors in the en vironm ent I 96

Regulat ion and interaction in Ecosys tem s

AL DO L EOPOLD--i94 1

L akes in relation to terrestrial life patterns 200

G. EVELYN HUTCHI NSON - I959

Homage to Sa nta R osalia or W hy are there so many kinds of anim als? 204

ROBERT H. M AC ARTHUR AND JOHN W . MAC ARTHUR-1961 On bird sp ecies div ersity 20 8

EU GEN E P . ODUM- I 962 R elation ships between structure and function in ecosys tems 2ii

RAMON M ARGALEF-1963

Oil certain un ifying principles in eco logy 215

INTRODUCTION

It has been said that eco logy is as diversified in its scope and mean ­ing as are the diverse Ho mo sapiens that call them selves ecologists . While th is belies the situation, it d oes intimate the great disparity in orientation and approach to the field. One of the benefi cial re­sults of this div ersity has been th e accu mulation of a wealth of data and interpretation providin g ferti le gro und for the develop ­ment of th eory . Among the less salutary consequences o f th is pro­tean posture has been the judgment that eco logy is but a point of view rather than a scientific discipline. An anthology provides an opportunity for different ecologists to react indirectly to that criticism and in so doing to indi cate the particular province of nature which they hav e carved alit as their special area of inquiry.

Underlying the varied form ulations describing eco logy is the the me th at it d eals with the interactions of organisms and environ ­ment. Although Haeckel is usua lly credited with the event, R eiter appears to have been the first to com bine the Greek words oikos ( = house) and logos (= study o f) to form the term ecology ; this was in 1 8 6 5 . It was H aeck el, however, who in 1 866 first gave definition to the term as "the body o f knowledge con cerning the

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