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TUNDRA AND
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TEMPERATE DECIDUOUS FOREST

GRASSLAND
DESERT
TROPICAL FOREST

TEMPERATE RAIN-FOREST

of

Me

of

Ecological

Animal Geography

Ecological

Animal Geography
SECOND EDITION
W.
'tul

C.

ALLEE

and

KARL

P.

SCHMIDT

Prnfcfmtir of ftwlugy

Chief Curator of Zoo'logy

llnirrrsity of llorida

Chicago Natural History Museu

AN AliTHOHI/KR K1HTION, KKNVBITTEN AND HEVISKD
i)

ON

'r\cr<!f'ogr<i)>hi(' aiif

oekologischer Grundlagc

By

the laic

RICHARD HESSE

NEW YORK
LONDON

JOHN WILEY & SONS,

INC.

CHAPMAN

& HALL, LIMITED

COI'YHICIIT,

1937,

1951

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UMVKHSITY

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CHICAGO

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Published

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Jolm Wiley & Sons, Inc.
Tlui Unixersity of Chicago Press

By arnni^cmcnt

vvitli

THE AMERICAN EDITION
is

dedicated to hco inspiring teachers:

THE LATE DAVID WORTH DENNIS
of Earlluim College

and

JAMES G.

NEEDHAM

Preface

to the

Second American Edition

logical

The preparation of a comprehensively revised edition of our EcoAnimal Geography of 1937 converts the work into a book still more independent of the German original. The 1937 text, though based on a close translation, had already been much revised by
changes, deletions, and additions from Richard Hesse's pioneering Tiergeographic auf oekologixcher Grundlagc of 1924. Even so, large blocks of the original remain, and we have preferred to retain Pro-

fessor Hesse's

name

in association

with ours, the more so as he himself

welcomed the idea

of an edition in English. When the junior author called on Professor Hesse in Berlin in 1932, the somewhat diffident

suggestion that considerable revision of the German text was planned for the American edition was received with wholehearted enthusiasm.

made by us would be scrutinized by himself German edition. The result of the incorporation of fourteen
further changes
4

Indeed, Professor Hesse remarked that any tiling added or any changes for a projected second
years'

accumulation of

and additions is an essentially new work. Those who to read the book in order to discover or interpret what wish might Hesse wrote or meant in 1924 must be warned that this is not the function of the work envisaged by us. We have hoped to make the book more readable, more accurate, and more nearly abreast with current knowledge; but we have frequently found Hesse's basic data and references of 1924, from much older literature, difficult to replace.

illustrate the nature and scope of our reand additions: aerial transport in distribution; bioluminescence in marine and cave animals; Bergmann's, Allen's, and Rensch's rules; the concept of the biome, with a schematic map showing the biomes of the* world; bipolarity in marine animals; coral reefs and islands; the Hawaiian fauna; intertidal zonation; the limnology of Lake Baikal and of Lakes Tanganyika and Nyasa; the Panamanian land bridge and the Tertiary Panama strait in relation to the marine faunae; the phase theory with regard to grasshoppers and swarming locusts; savanna,

The new material is thus and we make no apology. The following subjects

quite inextricably interwoven with the old,

visions

vii

viii

Preface to the Second American Edition

steppe, and desert communities; territoriulity; water masses in the oceans; winglessness in beetles; and "zigzag" adaptation. These items are not sharply set oft' either in importance or by length of treatment

from many other changes scattered throughout the book.
Revision of data concerning the gions has given particular difficulty.
of this character for

number
It is

of species in different rehard to find new information

surveyed our text.

may

and the regions more recently well have different boundaries from those discussed in

many

localities,

In specific instances,

when new and comparable data were

found, the significant proportions remained much the same as those derived from older lists, though there have been shifts toward both
increase and decrease with the changing evaluations of the species
lists whenever possible. When dates have available, comparisons frequently been introduced into the text so that the date of the reported survey meets

category.

We

have modernized such

we found no new
the eye.

Chapter bibliographies have been expanded to include pertinent publications. A few references were dropped or replaced. Some of the more obscure items and some long series of citations of data are retained by citing the original German edition, i.e., Hesse (1924). Much documentation has been preserved when the items bear early

new

elates.

Age alone

is

little

in fact, in the field of
cial

indication of the value of a given report; animal geography, older papers may have spe-

value when they accurately describe conditions of existence as they were seen years ago. Titles have been added to all bibliographic items, and the details of citation have been checked for accuracy as

The revised bibliographies are arranged alphabetiauthors in the hope that this will facilitate their use. The cally by main burden of amplification of the bibliographic references has been
far as possible.

borne by the junior revisor, and in this the aid of his colleagues, especially Dr. Fritz Haas, Robert F. Tnger, Marion Grey, and Robert Kanazawa, and of Alan Solem, is gratefully acknowledged. A few figures have been replaced and a few have been added. The
treatment of the phenomena of the ecological distribution of birds has

been changed

constructive sugDr. Austin gestions by J. J. Hickey of the University of Wisconsin. L. Rand, Curator of Birds at the? Chicago Natural History Museum,
in several places, often as a result of

has also been especially helpful in this connection. Statements about rnollusks and certain other invertebrates have been scrutinized by another colleague, Dr. Fritz Haas, Curator of Lower Invertebrates. Other colleagues of the junior author at the Chicago Natural History

Museum,

especially Colin C. Sanborn,

Loren

P.

Woods, Rupert

L.

The senior revisor's students in successive classes in animal geography at the University of Chicago have frequently rendered aid in pointing out obscurities and in making pertinent suggestions. secretary to the senior rcvisor. departmental artist. Illinois KARL P. now of Lahore. and that concerned our main emphasis in the final chapter from the long-estabmodern man with the environment and with other animals in Europe to the more recent and dramatic impact of the white man's civilization upon the American scene (see p. In the complicated preparation of the revised manuscript. 1951 . Alice. has been invaluable. Thomas Park. He Hesse's approval of our unsupervised revision of his offered only one major criticism. Dybas. Although the two books are concerned with different problems. by W. Emerson. we have retained our former emphasis. Antioch College Assistant at the Museum. ALLEE SCHMIDT June. After reconsideration. Margaret Schultze. and to edition of more mature scholars for their cordial reception of the first American what we still think is a great book. bore the brunt of the preparation of the index.Preface to the Second American Edition ix Wenzel. and Karl P. Pakistan. Gainesville. have been helpful in suggesting additional matter and correcting nomenclature. C. Florida W. We are thankful to the late Professor Richard Hesse. Schmidt (1949). although with added discussion. Bauer. Miss Patricia Grubbs. secretary of the Department of Zoology at the Museum. The present revision was carried forward in close connection with the closing stages of the preparation of the Principles of Animal Ecology. in charge of illustration in the department. Alfred E. to Englishreading college and university students throughout the world. Miss Laura Brodie. Dr. We especially appreci- ate Professor work. the aid of Miss Margaret J. in the final chapter and elsewhere. 651). Emerson has been consulted about termite distribution. Chicago. Orlando Park. Alfred E. Muzaffer Ahmad. as was the assistance of Mrs. of the world-wide need for conservation of natural resources. C. and material concerning the zoogeography of India was read critically by Dr. and Miss Margaret Bradbury. documented by shifting the lished interrelations of 1 citations to recent pertinent books. preoccupation with the formulation and documentation of ecological principles and with the recent literature in that field made it easier to revise and modernize Ecological Animal Geography. and Henry S.

.

Such treatment does show that the problems of ecological animal direction.Excerpts from the Original Preface When I finished my essay on "The Ecological Foundations of Animal Distribution" which appeared in Hettner's Geographische Zeitschrift ( 1913 ) I had thought to turn to other work. the solution is to I be sought. I offer this book to the friends of zoogeography. we need rather . For the first time an exposition of animal geography is presented which gives approximately equal space to the animal life of the sea. of fresh water. participated have resulted in the collection of a mass of material almost too great for mastery. treatment will stimulate further expediWe have had an oversupply of travel xi which yielded animal pelts and alcoholic material. after twelve years of steady work. and of land. hoping for a favorable reception. for example. Ecological animal geography is a young science. marine biological research stations. These sources have yielded results of practical importance for the promotion of fisheries as well as of significance for theoretical biology. Researches upon animal life in fresh water have also come into the foreground in all civilized countries. hope that this tionary researches in this field. and but relatively little space is given to presenting satisfactory solutions. which was not completely interrupted by twenty months of army service. In this new field the fundamental questions are yet to be formulated in order that a I hope this rich phase of biology may be opened for further work. but this material had fascinated me and I could not leave it. and the great expeditions for the exploration of marine life in which almost all civilized nations have is very extensive. There is an understandable demand for such treatment. and its presentation cannot result in so clear a picture as. but the material to be mastered before this is possible Since the founding of the Zoological Station at have been opened an increasingly large number of there Naples. through observation geography are capable of exact solution and indicates further in what and experimentation. the classical "Lectures on Comparative Anatomy" of Biitschli. So. it deals largely with problems which are taken up separately and arranged in order. book may be thought of as such an attempt.

since that would have lengthened the list unduly. 1924 R. Hartert's comprehensive Since the different parts of the book have been. Germany Late March. Because the use nomenclature zoological for of Latin names have frequently Inasmuch as the facts used must be drawn from a rich literature. followed Grobben's revision of the Textbook of Zoology of C. my knowledge has not always sufficed to a correct name. for in many cases. and the designation "C. In special cases I have followed other sources. so much the less since. Temperature is given in degrees centigrade. guarantee is still full of contradictions. for palaearctic birds.xii Excerpts from the Original Preface observations on the relations between i\pimals and their environment. Bonn. I have been almost frightened by the accumulation of animal are not given as fuunul lists in which the species are grouped with no other connection than that they come from the same locality. and I have restricted myself mainly to smnmarizatiqns and to books. All measures arid weights are given in the metric system. 1917). work." is usually omitted. names many species. Glaus (third edition. despite all reforms. I have tried to bring together the most useful sources and have appended a selected set of references to each chapter. and other data have been changed to this when necessary. They In the naming of species I have the formulation of general laws. I have not cited all references that have been used.. rather the species are given as examples for names in the text. Hesse common animals is bombastic" given the common name only.taken from work done at widely differing times and since there are different e. I . I have used E.g.

8. 7. and Means of Dispersal The Effect of Geographic Isolation Historical Zoogeography The Influence of Extent of Range Biotopes and Biocoenoses Barriers to Distribution 68 89 119 147 165 PART Introduction 10. The Problems and Relations of Ecological Animal Geography The Conditions of Existence for Animals The Effect of Environmental Selection on Animal Distribution Classification of 30 41 Animals According to the Most General Characteristics^ of the Environment ' 5. 24. 1- THE ECOLOGICAL FOUNDATIONS OF ZOOGEOGRAPHY 3 15 ' 2. (1 7. 3. 19. 25. Jig- and Desert Communities Animal Life of Swamps and of Shores Alpine Animals The Animal Life of Polar Regions Island Communities Subterranean Animal Life The Effect of Man on the Distribution of Other Animals Index xiii / 453 508 533 565 588 608 621 636 650 675 . The Biotic Divisions of the Ocean: the Benthal a The Biotic Divisions of the Ocean (Continued): the Pelagial The Abyssal Benthic and Pelagic Communities of the Sea Geographic Divisions of the Animal Communities of the Sea 3- f PART Ifi. THE DISTRIBUTION OF ANIMALS IN INLAND WATERS. A PHASE OF LIMNOLOGY Factors in Inland Waters in The Environmental Communities Communities Communities in 17. Ecological Factors of the 21.Contents PART 1. in 347 362 383 431 PART 4- THE DISTRIBUTION OF LAND ANIMALS Land and Their Effect 20. 12. The Chemistry of the Sea and Its Influence on Animal Life 13. Grassland 23. 26. Running Waters Standing Inland Waters Other Inland Waters 18. Forest Communities l on Animals 22. 9. 14. 2- THE DISTRIBUTION OF MARINE ANIMALS 179 185 203 217 272 300 326 Physical Conditions in the Ocean in Relation to Animal Life 11. 15. 4.

.

Part 1 The Ecological Foundations of Zoogeography .

.

from protozoans and coelenterates to vertebrates. and 10t * from place to place makes animals to a degree independent of their environment. though very unequally. have been included. since all groups of animals. has dealt mainly with the distribution of the vascular plants. of plants. or warmth in new localities. in this respect. enabling them to cross barriers with less difficulty. Zoogeography corresponds to phytogeography aaid with it forms the single science of biogeography.studied. phytogeography has been the subject of active research for a much longer period and has accordingly been much more intensively . in special fields. On the other hand. 14. and zoogeography. the majority of them are at least able to move towards to water. Modern work in plant ~'* geography continues in bothenvironment are much The capacity for motion historical Among more ecological-aspects. food. in zoogeographic studies. the relations with the total direct and obvious than among animals. rests upon a broader basis. The formation of spores or seeds. Many physiological problems have thus been in Superior numbers refer to the respective items end of each chapter. These divisions of biogeography are very unequally developed. effectively protected from unfavorable influences and easily distributed passively. so that limitations of the distribution of plants in accordance with their ancestral history are very much less evident favors the wide distribution than among animals. See p.-' plants. the bibliographies at the 3 . Phytogeography. in general. The Problems and Relations of Ecological Animal Geography ZOOGEOGRAPHY IS THE SCIENTIFIC STUDY OF ANIMAL LIFE WITH REFERence to the distribution of animals on the earth and the mutual influence of environment and animals upon each other. and thus they become exposed new conditions. This branch of science therefore forms a department of both zoology and geography.JL.

vegetation. are enumerated for specified areas of varying extent. constitute another main problems for faunal zoogeography. Various habitats. Important scientific expeditions have brought and together great numbers of museum specimens from distant countries The record. Extraordinary advances have been made in this direction during the last century. Zoogeography trolling factors is made difficult and by the great by the great complexity of the condiversity of reaction to them among the varied types of animals. or ranges. beetles and butterflies. in which the animal populations. This is the function of "faunal zoogeography. defined biotopes and their dominant species or "indicators. or earthworms. according to soil. conspicuous forms. each inhabited by a definite and The determination of wellwell-characterized animal community. Animal life is very unequally distributed in any considerable area. springtails." plus the intensive study of the associated animals. of the individual species must be exactly series of . and snails and mussels. forms the basis for every further development of the subject." which goes hand in hand with the identification of animal species and in general with These studies are not equally adthe accumulation of collections. The solution of zoogeographic problems is not therefore less attractive or important. which will require much further study. mayflies. In addition to determining the species of animals for a given region. vanced in the various groups of animals. and their solutions more Zoogeographers have been able to examine many easily attained.4 more Problems and Relations clearly defined for the plant geographer. can be distinguished. such as birds and mammals. in identifying animals and putting in order a wealth of data. Faunal zoogeography. phases of animal distribution in the light of phytogeographic studies. The gradual development of the study of animal geography makes it possible to distinguish a number of diverse lines of research that appeared one by one with the advance of this department of science. or diversity of form. Classes and orders whose representatives are notable for their beauty. brilliant coloration. or whose simplicity of preservation encourages collecting. even in the most highly civilized countries. and climatic conditions. have always been more intensively collected and studied than such groups as the hydroids. and these have been described and classified. is far from complete for the smaller. squids. whether for single groups or in toto. seas. the specific areas. or biotopes. less however. The origins and foundations of zoogeography lie in the accumulation of faunal lists.

for example. but did not go so far west as the rhinoceros.Problems and Relations defined. among song rnelodia and Melospiza melodia sparrows. 1 not. The principal avenue of approach to faunistic zoogeography tematic zoology. made at the bordering limits of geographic ranges. This is furnished by the trinary system of nomenclature. the yellow Subspecies of such species may be sharply also be united by intermediates so that the able. the song sparrow. and antelopes are found together in the steppes of central Africa. The eland extends beyond the Zambezi-Congo divide. Of prime importance is the recognition of the fact that many widely distributed animals have a different ap- pearance in the different parts of their range and are divisible into geographic subspecies by means of constant minor differences. and the work of the systematists in this field. The black rhinoceros originally ranged to the extreme south of Africa. is sys- Intensive systematic studies have unearthed new problems for zoogeography. especially for birds and mammals. Many detailed studies of this type have been made. zebras. and exact knowledge of these is required to complete the foundation for zoogeographic It is evident that crucial ecological studies must be investigation. of this are lizard Lacerta the wall muralis. and. It is. but apparently never occurred south of the Orange. extending thence to Senegambia. rhinoceroses. most of the savanna mammals do Carncroons. Zebras also formerly ranged to the Cape. swallowtail Papilio tttnws. but they may practiced eye of a specialist is sometimes required to distinguish them. the wild turkey. to be desired that a uniform nomenclature and a definite characterization of these geographic subspecies be introtluced. The giraffe ranges from the Orange River to the Zambezi. whereas the zebra is present. the puma. A vast Buffaloes are absent in Somaliland. the of their general agreement in 11 12 * king snake Lampropeltis gctulus. is especially valuof course. and it extends no farther west than the Niger. in spite Some comimportant characters. defined. The recognition of these minor geographic differentiations is of importance in the study of the influence of external conditions upon animals. Its southward range is now much restricted. among butterflies. lions in the amount of detailed study remains to be done on the definition of the ranges of individual species. It is absent in part of Mozambique and only reappears to the north of Rovuma. type prehensive species the African lion. the listing of Melospiza melodia atlantica makes it immediately evident . 5 The ranges of species that occur together in a given locality may be entirely unlike. The giraffe appears to be absent in British Central Africa. Giraffes. although frequently regarded as trivial.

There is interest and even fascination in the recording of observations. an* distinguished in comparative anatomy. insects. The data accumulated by faunal zoogeography must now be sifted and ordered. for forms (now species) that remain dis- tinct in spite of overlapping distribution. preserves the useful trinomial grouping. Building materiel! is necessary to erect a building. who sentative. introduces the useful terms allopatric. faunae may be distinguished whose distribution does not agree with the present geographic divisions of the earth. The fauna of southern Asia is more reptiles. Comparative zoogeography attempts according to their resemblances. various directions. and Hornologies. but a heap of bricks is not a structure. When different faunal lists are compared with reference to the natural relations of their components. and their interrelations studied. or acquired resemblances. leads directly to classification of the facts of distribution. The natural laws that lie hidden must be established. Many groups of it is to the fauna North American birds differ more widely from the corresponding groups and South America than from those in Europe and northern Asia. The definition of subspecies as geographically representative forms." The term Formenkreis has become current to distinguish series of allied forms whose distinction is geographic and whose origin may be supposed to be entirely or primarily accounted for by geographic isolation. The most obvious faunal boundary between the animal life of North America and that of South America lies somewhere in Mexico and not at the dramatic Isthmus of Panama. but only the hod carriers of science could be content with the mere accumulation Faunal zoogeography. regardless of contiguity of range or of intergradation. and sympatric. The homologies among mammals and in Central . to classify animal distributions This comparison or may be made from resemblances. and an accumulation of unrelated basic data is not a science. it brings the definition of species The definitions are congruence. therefore. this classification may proceed in of facts. closely related to that of trans-Saharan Africa than of Asia north of the Himalayas. Such animals of North Africa as the snails. for forms not found in the same territory and thus geographically repreand Formenkreis into 9 essential sharpened by Mayr. birds. and a similar distinction applies to animal distributions. In fact. different viewpoints.6 Problems and Relations that the forms mentioned belong to a "Formenkreis. and amphibians are much more closely related to the aniof the mals corresponding groups in southern Europe than to those of Africa south of the Sahara. inherited analogies.

while flying animals are Water animals are limited in their and land animals by water. The position of the and approach more important bar- with geologic changes in the earth's occupy much shorter periods of time than are required for changes in the means of dispersal through organic evolution. We know that many places now occupied by land were formerly covered by seas. spread by land. uniformity within the group. may be separated. and regions may be united whose faunae were only distantly allied. in different natural groups of animals. unable to pass mountain ranges. We to the animal. they are as old as the principal subdivisions of the animal king- dom. . are interrelated and of common origin whether they inhabit the forests. within which the animal inhabitants are homologously comparable. and these alterations America and Eurasia (via Alaska). that areas formerly well watered may now be desert. which were repopulated after the withdrawal of the ice. Climate. the external barriers affect the dition the distribution of a species: the and the different groups in diverse ways. It is highly probable that land connections formerly existed between certain regions now separated. The means of dispersal remain unchanged through long periods of time. formerly continuous. that high mountains may be raised anew while others are eroded away. and North however. insects. the ovenbird family. such as the echinoderms. For example. such as North Africa and southern Europe. or the presence of more successful competitors or of enemies may present barriers to the dispersal of any group. or the mountains. and the rodents. Closely related species of animals will in general have adjacent ranges. and that ice sheets extended over previously inhabited regions. means of dispersal available or biotic barriers to such disexisting physical As a of the differences in the means of dispersal consequence persal. since it is to be assumed that the area in which they devel- oped from their common ancestors was the common origin of their observe in this respect that related human stocks in Two factors mutually congeneral have a continuous distribution. alters surface.Problems and Relations 7 such comparable faunae are based upon the blood relationship of their components and upon a common evolution in time and space. The larger faunae of this kind characterize the faunal regions and their subdivisions. such as the iguanid lizards. that rivers had other courses in former times. the lack of suitable food. the prairies. riers to dispersal. Land animals are often least affected by barriers of any kind. fishes. the representatives of natural groups in South America. distribution. or birds. Through changes of this nature the ranges of related animals.

and the horseshoe crabs on the east coast of North America and in the Moluccas. tion were: (1) via Java. and the more opportunities for dispersal will have been available to its members. or tardigrades. In many instances. America and in Malaysia. antedating the changes in distribution. from the by communities of animals dormant state for long periods. South Africa. peripatus in New Zealand. nematode worms. exhibit a whole series which adaptations for climbing and among for parachute jumping are especially notable. hummingbirds. rotifers. Africa. tropics to the polar regions. matter will then consist of such problems as the restriction of like the penguins. resemblances in their possession of adhesive apparatus. and South America. lemurs. which entered Celebes at succes- These highways of immigrasive periods over four distinct routes. Beds of moss. of evident resemblances. South America. such as the tapirs in tropical . and the fauna of a given region may be analyzed by studying the distributions of the subordinate faunae of diverse origin represented in the area. for ex14 Four ample. and (4) via the Moluccas. For example. Asiatic and Australian elements are intermingled. or armadillos to specific areas.8 Problems and Relations The older a natural division of the animal kingdom. The animals in mountain streams in all parts of the world have numerous and surprising countries. (3) via the Philippines. are inhabited characterized by the capacity to live in a . Historical zoogeography in this way attempts to work out the development in geologic time of present-day distribution by studying the homologies of animal distribution. exhibit numerous resemblances in their composition. at considerable distances from the continents. the more such changes of barriers will have occurred during its history. the absence of otherwise widely distributed ject groups forms from certain particular areas (as bears in Africa south of the Sahara or of placental mammals in Australia ) and the presence of related forms in widely separated regions. (2) via Flores. the inhabitants of the rain-forests of the various tropical and Malaysia. copeThese biocoenoses found in a given habitat pods. On the other hand the geographic unit may be taken as the starting point. monotremes. 1 -' 1 - immigrations can be distinguished. In Celebes. their environments. Ecological communities of animals may be recognized that resemble each other superficially in correspondence with resemblances between These are analogous rather than homologous. The faunae of small islands. whether they be protozoans. For such studies the starting The subpoints may be the systematic groups of related animals. the systematic relations are the primary factors.

but on account of the differences in their methods the relative value of their conclusions is very unequal. reasons for It is mals. This phase of zoogeography may also proceed either from a geographic viewpoint or from the animal itself. omy" of plants and animals. As in human history. In this way an explanation may be found for the convergent evolution of different animals under the influence of similar environmental conditions. Ecology is the science of the relation of organisms to their surroundings. The of approach deals primarily with the geological history and with the phylogeny of the animal kingdom. as contrasted with the historical. or the ecological relations be- tween an environment and its animal population. 9 in the different faunal The ecological viewpoint. the peculiarities resulting from the influence of the surroundings. selective operation of habitat requirements on the composition of the fauna. whether in America or Africa. the modifications in appearance and habits underby the animal population in adaptation to the given conditions. lems is sought by causal zoogeography. in their it is the science of the "domestic econ- Ecological zoogeography views animals on the of their native regions. location of this region. the events of geologic history and of animal evolution historic mode of the earth .Problems and Relations have analogous communities in similar habitats regions. in their conditions dependence to their without reference to the geographic adaptation surroundings. the northern or the southern hemisphere. If a specified animal is made the starting point be pursued. regards the analogies between animal communities in similar habitats. The answer to these prob- phenomena. rable only a step from the observation of all such groupings of aniand from the recognition of the fact that they involve compa- to the questions of the causes of the appearance of with varying limits or of the causes that condition the chargroups acteristics in groups with similar habitats. living as well as non-living. The geographic questions concern the requirements created by the environmental conditions of a given area for the structure and habits of its gone and the inhabitants. the questions \yill concern the anatomand physiological characters that fit it to its surroundings and enable it to compete successfully in the struggle for existence. to be studied are According as the associations homologous or analogous. causal zoogeography studies the historical reasons for the evident differences in distribution of the natural groups of animals. and the for the studies to ical its failure to spread into other environments. The results of historic and ecologic studies in zoogeography are mutually supplementary.

The reconstruction of such past events is consequently uncertain. It is true that the position occupied by an animal is conditioned by its heritage. In an important work on the polyphyletic origin of Burckhardt tion. and the number of erThe abundance of incomroneous conclusions excessively large. It is zoogeography accordingly one of the most important problems of ecological to investigate the adaptations of animals to their sur- roundings. it studies processes.e. and the modifications thus acquired are frequently of an adaptive nature. the thickening of the under the influence of wave action. i. but environment also influences the ani- The mal. ecological method is quite different. and it has been fruitless to seek in them for universal causal connections. The enlargement of the often mammalian ment shells of mollusks heart with increased bodily activity. as was attempted by Reibisch and Simroth with their pendulation theory of climate and by Eimer with his orthogenetic theory of species formation.. Examples are the reduction in size of marine animals with decrease in the salinity of the water. Ecology deals with present conditions and phenomena. Such directly conditioned changes may accidentally prove .10 Problems and Relations are never twice exactly alike. in another direc* by false phylogenetic premises. and the enlarge- for excretion are examples. by natural selection if by no other means. which can be repeated. Such processes are necessarily repeated when the same conditions are supplied. Instead of being concerned with unique events. between continents has for connections former land patible hypotheses been shown graphically by Handlirsch. adapWhether an organ is passive or directly active. An animal may become adapted to the conditions of its The most frequent existence by somatic and by genetic processes. and most important form of adaptation is somatic. has brought out the mistakes produced. 5 who figures all the supposed land bridges of Cretaceous and Tertiary time on the same maps. owing to the wonderful property of living matter to react adaptively. tation. such as chemical reactions or physical experiments. of the kidneys in fresh-water animals with increased necessity Other somatic changes that appear with equal certainty in consequence of environmental stimuli may be of indifferent value to the organism. scarcely a bit of ocean has escaped the supposition of having formerly been occupied by land. the large terrestrial birds of the southern hemisphere. they make it easier for the animal to live in its environment. which are subject to analysis and repeated test. its capacity is increased by use. or functional. and many changes in coloration induced by increased or decreased tem- perature.

Artemia salina. Experimental animal undergo active development. to be sure.Problems and Relations to 11 be of value to the animal. in the small crustacean of saline waters. and of the relations between frogs and their water intake. and they may then be designated as coincident adaptations. and thereby cause the animals to resemble the desert floor. and zoologists are less advanced than the botanists is in this field. where they were absent. Field experiof also turbellarians ments have yielded important results. still at the beginning of an experimental ecology. of the transformation of the salt-tolerant crustacean Artemia salina with changes in the salinity of the water.. although present in the mainland brooks. Many processes of this kind can be verified experimentally and may be made the subject of physiological analysis. change of direction new adaptive radia- Such "zigzag" evolution is exemplified by the derivation of the rays and skates from the sharks. In this way the colors of the most diverse animals lose their brilliance under the dry heat of the desert. The darkened coloration exhibited by many Lepidoptera when subjected to cold during pupation may favor the warming of the body of the adult in the northern species. species have been introduced into the brooks of the island of Riigen. are the more neces- water. We may call attention to the kind of preadaptation that involves of adaptation and may afford opportunity for a tion. The study of adaptations is for this reason of especial importance to an understanding of the ecology of animal distribution. supporting ability) of the salt solution. sary to animals the further the conditions of their environment depart from the optimum. become pale and dull. physiological or structural. on account of the abundance of interesting results promised.e. These mutations are for the most part of no importance to the efficiency of the animal. 16 Special adaptations. Again. Germinal changes also give rise to new characters in animals. the flattened plane of the belly of the free-swimming shark. Laboratory studies have been made of the change of form in daphnias under the influence of food and temperature. We are. a Danish race of Daphnia cncuUata was introduced into Lake Nemi in the ecology certain to . but they may sometimes be of value. which aids it in maintaining position in the is evidently the preadaptation that governs the mode of further flattening in association with bottom-dwelling habits. and they may be preserved by selection and become more and more widely distributed and then more and more adaptive. the relative size of the supporting surfaces is increased with decreasing density (i.

a sound body of historical zo- ogeography is growing up as paleozoological knowledge increases and is brought into harmony with faunal zoogeography. Nevertheless. remarkable that one is able to find clues to events of the remote of historical its The aims valid answers to Hispast by the analysis of the homologies in animal distribution. problems.* 7 * The introduction of English foxes into Australia and of the muskrat into Europe are further examples from the very long list of such natural experiments. . and It is questions would be of great importance. The numerous foreign insect pests brought into the United States have been much studied. when the study of homologies overshadowed all other linos of research tion and the causes During that period. torical zoogeography has been valued highly in the eyes of numerous investigators because of its endeavor "to unravel the history of the colonization of continents. was actively investigated. it must be admitted that more sins were committed in this division of zoogeography than in any other through the proposal of unwarranted and frivolous hypotheses. The whitefish introduced into Lake Laach (near Coblenz) have transformed into a new subspecies (Coregonus fera bcnedieti). The data are often meager. and their results throw light on certain questions of ecological zoogeography. The release of the English sparrow and of the European starling into North America (in part by the same person!) are familiar American examples. 1 perimental stud\ of the problem of ecological zoogeography gives to the results obtained a potential degree of certainty quite in contrast with the frequent uncertainty of the conclusions of historical zoogeogr raphy. special races of Peridinia and of Anodonta have been introduced into newly made artificial ponds in order to study the ensuing changes. such as that of the permanence or transitory nature of continents and oceans. On the other hand. Unplanned experiments have been still more numerous. zoogeography are unquestionably high. and the sources of error great. geoNo unilogical evidence wanting. A most illuminating and comprehensive work on this subject is Thomson's The Naturalization The possibility of an exof Animals and Plants in New Zealand. historic zoogeography was enthroned. and to discover the highways of distribuof migration in past epochs. of in has been reached even the most fundamental opinion formity in zoology.12 Problems and Relations Alban Hills near Rome." 14 This was particularly true in the half century following Darwin's Origin of Species. and now exhibits in consequence a fine series of well-established and connected results.

Experiments with populations. it has established some general as the application of the law of minima to the phenomena such laws. as follows: A. Paleontological. d. 1.Problems and Relations 13 In contrast with the speculative nature of much historical zoogeography. mostly with new emphases limnology. 20 Zoogeographers have by no means been in full accord. Comparative a. b. Experiments on tolerance. 3. with . Ecological zoogeography. accumulating data from field studies. It must be pointed out that a major divergence in speculations and conclusions has occurred between historical animal geography and historical plant geography. Phytogeographers have sought explanations for large-scale distributional phenomena in east-west or west- east connections between the continents in past geological ages and have postulated the origin in the southern hemisphere of great groups of plants that had subsequent northward dispersal. of distribution (cf. and causal. Chapter 2). 3. s>15 hemisphere by existing Phytogeog- Zoraphers have freely invoked land bridges or continental drift. searching for analogies in the data thus found. Historical zoogeography. . we arrange the subdivisions of zoogeography. Experimental. a by-product of systematics. 4. field studies. have mainly postulated is reviewed by Just. and science. 2. stability of the continents. c. searching for homologies in the data of faunal zoogeography. Test of theoretical conclusions. the correlation between the weight of the heart and the isotherms of Further active research will produce a blossoming of this science like that of its elder sister. land-bridge speculation and after a thorough review of the evidence from fossil mammals. climate. accumulating data from ( B. Experimental. oceanography 1 ) . Faunal. tracing existing distributions into the past. ecological zoogeography bears the germs of a more truly causal Although still in its infancy. 1. in the past generation. but they tend to search for the origins of groups of land animals in the northern hemisphere and 1 '*' to account for their dispersal to the southern or known former connections between the continents. Botanical opinion By way descriptive of review. ecological phytogeography. 2. Fuunal. Comparative. Bergmann's Rule (Chapter 20). examining capacities for dispersal. Examination of introductions. after a of uncontrolled period ogeographers.

27 figs." Ibid. Lcip/ig. 223. "Geology and plant distribution. W. "Die Siisswasscr-Mollusken \-n\i Celebes. S/. 1943. Marion I." Ecol.. Figs. Bernhard. Columbia: xiv." 4. Syst. Handlirsch. "Die Felt-hen des Laacher Sees. "Das Problem des antarktischen Sehopfungseentrums vom Standpunkt dor Ornithologie. Chronica Botan. "Beitrage zur exacten Biologic. 15 pis. G. Maemillan: Klages. 1915.stetnati. 17:159-183.. Systematics and the origin of species from the viewpoint New York. 1913. 1 8. London.. 1 map. Sitzbcr. 5 maps. Stanley A. . 5. 1901. 63 figs. D.-n.. P. The naturalization of animals and plants in Ncu: Zealand. Engell. 169. B. Kurze Antceisung fiir ziwlogisch-sy. Foundations of plant geography. 2 maps.. 1898.sche Studicn. Newbigin. August. Univ.. Plant and animal geography.. xviii. 615. Monthly. Thienemann. H. Ernst. 6. l:\iii.." Sri. George M." Ann. 7. Rudolf. 12. 33 figs.. Materialien Naturg." Zool. of ancient dispersals. Cambridge.14 Problems and Relations BIBLIOGRAPHY 1.' Am. 1943. 1944. 17:127-137. Monographs.. Sri 24:171-318. 116. F. 18.. "Distribution patterns in modern plants and the problems Ecol. W. 108 figs. 1947. 171:1-84.. 30:241-253. Just. Akademische Verlagsgesellschaft: iv. 1):361-481. Simpson. 1929. 298. Problem der Artbildung. 1934. Harper: xiv. 32:173-220. "Die Silbcrfelchen des Laacher Sees. 1-32. M. v).. Erganz. G.. 22 figs. Acad. Methiien: xv. 607. 206. C. Schmidt. "Corollary and commentary for 'Climate and Kxolu14. 64:481-495. 1947. 20. xv. "Verbreitung und Haufigkcit des Elefanten nnd Loweii in Afrika." Jahrb. New York.. H. Berlin." Petermanns Mitt. and C. 16. 15.vf. 1936). Idem. Anton. 1947. 35 figs. Jahrb. Theodore. Idem. 2. W. Bemerkungen zu dcm glcidtbetitelten Aufsatz Victor Bauers. ( 1st ed. Y. text figs. Ecological crop geography. Camp. of a zoologist. K. No. 17. "Climate and evolution. Zool. Midland Naturalist. " tion. N. Press: x. Wien (m... New York. 9. "The problem of plan and purpose in nature. 3: vi." Zool. 3. Waltham. Cain. 9 19. 1942. Thomson.. Idem. "Uber die geologische Geschichte der Insel Celebes auf Gnu id der Thierx erbreitung.) 122 (Abt. K. Anz. An introduction to historical plant geography. 556. Borntraeger: 1. 1922. 1 map. Mayr. 1912. Das Princip geographischer Rasscnkreise itnd das . 1948. 104. E. 29 figs. Celebes. 75:226-234. 1902." Wiss. Sarasin. text figs. Matthew. 15:499-536. 11. 1911. 1942. 334. Wulff. Burckhardt. V.. 1928. Akad. 39 figs. 36.. 13 pis. Monographs. 10.. P. Rensch. Sarasin." (p. 13.

protozoa. blue- given algae. animals are unable to manufacture food from inorganic substances. 15 . or in fumaroles. for no animal is able to exist in its waters. in fact. and large numbers of the brine shrimp Artcmia ( Artemisia) and of the brine fly Ephydra. sodium. and and some of these may be imbibed independently of food. few localities ing to environmental conditions. 3 Animal life is thus very widely distributed. The Dead Sea merits its name. LIVING ORGANISMS ARE FOUND everywhere on the earth's surface in places to which they have access and in which they find the necessary conditions for existence. calcium. unlike green plants. By contrast. One seeks in vain for animal organisms in springs rich in carbon dioxide. and they are exceedingly rare in the ice wastes near Sea. among others. which is released as work and heat in the body of the animal. with a salt content almost as great. The depths of the Black which contain much hydrogen sulphide. This energy is developed to best advantage in the presence of oxygen. For animals these are primarily organic matter of animal or vegetable origin since. see Chapter 19 ) and in the depths of moving sand.The Conditions of Existence for Animals OWING TO THEIR RAPID REPRODUCTION. contains diatoms. Organic nutriment supplies large amounts of chemical energy. Great Salt Lake. supply of matter and energy is necessary for the existence of The substances required for the growth and reproduction of an ( organism. the poles A life. aside from the craters of active volcanoes and recent lava flows where then* is no animal life and where life is probably impossible. but its abundance and diversity vary greatly from place to place and from season to season accordThere are. Special salts supply needed elements such as potassium. are devoid of life. and for the production of secretions are designated as foods. for the maintenance of its energy. with their great concentration of dissolved salts and the high concentration of chlorides and bromides of magnesium.

potassium. It is true that many animals are capable of existing with a very small water supply and are extremely resistant to desiccation. magnesium. Protoplasm is full of water and suspends its functions when this water is withdrawn.the water. These condifies tions are closely 13 29 The investigations of approached by sea water. The larvae of Calcispongia and of sea urchins are unable to develop a skeleton in water free from calcium. and other simple types of animals retain their vitality in spite of long-continued drying even in sunlight. and other materials. The^olubility of inorganic substances in water satis- one of the primary requirements for the development of life. Tardigrades. It diffuses into their cells and dilutes the salt solutions of the protoplasm to an excessive degree. in solution. although animals also obtain many salts directly from . and of plant life especially. calcium. for more than three weeks. Some of these. 1 3 ' 5 ' 15 '" 4 ' 1 1 - by the chemical Meal worms (larvae of the beetle Tenebrio molitor] have been known to live in bran dried at 105 C. whereas the animals at the beginning of the experiment contained only 61. and as the chemical transformations in question can take place only in solu- tions. is the most favorable medium for animal life. water. The survivors of this experiment contained 65% water. Pure water is harmful to organisms. as a solvent.5? water. Sufficiency of water is most completely assured to animals that live it. is Some energy and heat. others secure water so-called water of metabolism ' transformation of their food. i. and become active with renewed water supply (Chapter 18). they usually contain. and some have survived the fourth week.e. form of light Water Water cannot be withheld from living organisms for continued periods of time without injury and ultimate destruction.. The waters that collect on the earth's surface are rarely even in approximately pure. water isotonic with protoplasm. whereas . The release of such energy occurs by means of chemical transformations. obtain water from their food.16 Conditions of Existence which is important for its own wealth of chemical energy. rotifers. greater or less amounts of the salts of sodium. both herbivores and carnivores. is also directly available to animals in the an indispensable requirement for life. and others have shown that sea water is approximately Fredericq isotonic with the body fluids of the marine invertebrates. Water containing salts in such concentration that no osm6sis takes place between it and the organism. nematodes.

and no important animal phylum is The circumstance that only marine organisms are known from _the older fossiliferous geologic strata also speaks for the marine origin of life. the ocean. whereas important groups are wholly absent on land and iu fresh water.7 per cent salt in solution. most snails. whereas. Nereis divcrsicolor and among The amphipod Niphargus occurs the salmon and stickleback.* restricted to these environments. . arc influenced by slight changes in salt content. that occur elsewhere must have de- veloped from marine ancestors and freed themselves from the marine habitat in the course of geologic time^ This hypothesis is supported by the fact that almost all the main branches of the animal kingdom are . 10 . for example. * limits between which animal limit is The lower is is possible defined by the necessarily activity The opposite theory maintained by some authors. The ocean accordingly affords its inhabitants the most favorable conditions of life*_a circumstance that has been explained by the supposition that life originated in sea water v ^ According to this theory all non-marine organisms.Temperature 17 thisjs_not true of fresh water and its inhabitants. Humidity .especially Euryhygric animals are numerous in the insects^ birds. as.represented in. be salt content in designated as stenohaline. in that withstand great changes as eurylialine. Temperature The temperature are not very wide. others withstand much variation in this respect. but some occur only under very humid conditions. and those may changes fishes. whereas others are partial dry situations. as is well known. amphibians. Salinity The different for different animals. water with 8. and water buffaloes are examples of stenohygric animals in moist situations. Worms. the brine shrimps of the genus Artcmia 15 The animals affected by slight tolerate great changes in salinity. the annelid . such as reef corals and am- phibians. Animals tied to narrow limits of variation in atmospheric humidity may be referred to as stenohygric.. and many groups. among mammals. .. and those that can withstand great variations of humidity are curyhygrif.Terrestrial animals cannot dispense entirely with water. extent of the variation in salinity that can be borne is very Some. the camel is a stenohygric animal to of arid country. both plant and animal.

minimum. 1 '- For individual ather narrow.) for the eggs of trout.or heat-tolerant.5 When and 24). when narrow.other hand. effects of the The two temperature limits are widely different. ranging from Canada to Patagonia. . The range species. (5^J^J^th.body . temperar tures usually causes the death of the animal^. the more liquid fats and Raper - 1 made solid occur in animals adapted to lower temperatures.of . or the tiger. and the large predatory cats such as the puma. minimum nearly C. the suspension of activity as a result of sarily fatal. J2^ezingjeiiipiature of tha . Stenothermul animals in turn may be cold. The tem- peratures tolerated by desert insects and desert lizards correspond 51 closely to those endured by the animals of hot springs. but active Protozoa tolerate only about 50 C. the oyster = Galba truncatula). the snail Limnaea truncation Germany is found principally in cold in the Pyrenees. (maximum 12-15 C. The upper limit of temperature is approached and sometimes exceeded JnJKbf springs. and 50 C. and 38-39 to the temperature range for a species is wide. between 14 and 20 for the eggs of carp. the temperature range is for the most part varies for different species but is fairly constant for a given form. about 55 is also the upper limit. the animal is said be eunjthermal.fluids. _ The suspension of the activity. mpjiLact^ ing matterjies^between 40 C. JThe iipper limiLoLtemBfirature. and the more fats are found in animals adapted to higher temperatures. a few degrees below that of pure water. The optimum may be widely different for different animals. The three cardinal temperatures are the naximum. ' the suggestion that temperature toleration is a matter of the relation of fats rather than of proteins. f Qj. the sperm whale Physeter. on the. is not necesmay be resumed with the return of a favorable temperature. (maximum 30. Encysted forms are more resistant. at whiclLpoiat-thcL^^ albuminoids jrgh6h!y. the optimum temperature usually ies nearer the maximum than the minimum limiting temperature. Many animals are accordingly able to exist in regions where they are forced to suspend their activities at certain seasons on account of low temperatures. which ( springs but lives in (2 to in 20). warm springs at 40 . about 22 at for frog's eggs for fowl's eggs. for Metazoa.Leathes .higlxes_t reported by 8 Brues). ). it is stenothermal. and active life low temperatures.protoplasm piodiiced by~-kigh. it lies between 1 and 4 C.. and optimum.-!Tl^f!I!gO d^tmr:tlMfi.jrJiPir>1 f^l ^hang*. . Examples of eurythermal animals are the flatworm Planaria gonocephala (limits +0. which lives in all oceans.18 Conditions of Existence .

in clays M 15. per kilogram and day. the salpids. 104 cc. 300 cc.5 12 9. latitude. Stenothermal warmth-tolerant animals are repre- sented by the reef corals. at 9 C. also increases with in- grcasiiig^tcnaperature^ The meal worm pupa uses. The eggs of the rotifer Notomma hatch in 4 days at 15. for example.-This is especially striking in the developmental stages.5 and at 16.5 and inus. the crustacean Calanns finmarchicus.5 Period of development..> 10 10. and 529 cc. the carp (which require at least 18 to breed). the mountains of Central Asia up to 4000 m. The rate of development of the eggs of the sea urchins Sphaerechinus and Echbetween 2.7 14 8.7. the crustacean Copilia mirabilis (between 23 and 29). divides once in 24 hours at 14-16. figures apply to the development of the cod Gadm Temperature. and 1692 cc. in 2 days at 28. and many others. and whitefish.5 <i 8 K>. Cold-tolerant stenothermal forms are represented by a cave silphid beetle that lives in ice grottoes where the annual 1.r> 5 17. . on temperature. The rate of reproduction for protozoans follows a similar rule: Paramccium aitrclia. measured by the consumption of aiuTtTTe production of carbon dioxide. at 18. The carp.0C. at 32.7 to +1. " The period of pupation in insects similarly depends twice at 18-20 1 . life Increase of temperature to a certain point reacts favorably on the of an animal because the chemical reactions on which the re- depends are accelerated by the rise in temperature... of oxygen at 15 C. and as far north as Irkutsk at 53 N. but require 40-50 days at the lower temperature and 6-8 at the higher. the meal worm Tcncbrio rnohtor requires the following periods from pupation to transforma- According to At (\ i:u 11 Hi 17 59IJ *r IV20 ^7 17^2 :w Kit Hours The oxvgen rate of animal metabolism. is increased 2. 15 the pearl mussel temperature ranges from and most reptiles.. tion: Krogh. which flourish only at temperatures above 20. Margaritifcra trout margaritifera. uses 661 cc. C -1 4^2 +3 4 '20.5.Temperature which is 19 equally at home in the tropical jungles of India. per kilogram and hour. the termites. at 25..2.5 times for every rise of 10 lease of energy 25. :J - The following callarias.5 The eggs of the herring develop equally well at +0.

The most widely known is that of Van't Hoff. as required by Van't Hoff. L>5 Ludwig reviews the literature and cites original observations on the rate of Popillia japonica development of the egg and pupa of the Japanese beetle which show that the temperature coefficient tends temperature increases. - temperature curve is a straight line that crosses the temperature axis near the threshold temperature of development. based upon changes facts in many 1 temperature. within these limits. Within medial temperatures thermal constants of development can be calculated by multiplying the number of degrees above the threshthe old temperature by the number of hours or days required for the development of the stage in question. Krogh's formula is based on the observation that within normal temperature limits an increase of 1 has as great effect upon metabolic processes at one temperature as at another. which is based on the observation that chemical and biological processes.20 Conditions of Existence Several formulae have been devised to interpret these temperature 3 effects. still widely used in biogeographic studies in America despite well- known defects. The life zones of Merriam. 44 The relations between temperature and rainfall or temperature and humidity for a given area can be shown by plotting the monthly mean by the rate of air temperatures against relative mean monthly rainfall or against mean monthly charts in humidity. Figure 1 shows two such temperature-rainfall which the vertical axis gives temperatures in degrees Fahren- . an hyperbola. within favorable temperature limits. and the physiological action of both by the is ac- affected movement. Many experiments show that such relationships hold for medial temperatures but that the rate of at development is greater than that called for by this formula low temperatures near the threshold of development and less near maximum temperature tolerated. The timetemperature curve is. in absolute to decrease regularly as the The Arrhenius formula. better fits the instances that show such a variation in the Van't Hoff Krogh-' advanced a formula to express the relation between temperature and the rate of development in which the constant is added to the rate at one temperature to obtain the rate at a higher one rather than multiplying the slower rate by a constant. are available in different altitudinal or latitudinal zones/' based supposedly upon the number of day-degrees The tempera1 ture relations of terrestrial animals are affected markedly companying humidity. are increased by an approximate constant (two to three times usually) for each 10 rise in temperature. and the ratecoefficient.

on data from Barro Colorado Island in the Canal Zone in Panama. scale.11 65 1833 60 15.56 55 101 12. Temperature is given on the vertical.7 80 75 23.Temperature and Rainfall heit or Centigrade. This is based in centimeters.89 20 1 23 4 567 6.89 70 21. The lower graph extends along monthly changes in rainfall.67 8 9 10 11 12 13 14 15 16 17 FIG. 1. and rainfall on the horizontal.67 30 1.11 25 3. and The graph at the F c 26. it the temperature axis with only slight summarizes these two elements of the climate in Chicago. Graphs showing mean monthly rainfall and temperature for a temperate climate (lower).78 50 10.22 40 4. and for a tropical climate with wet and dry seasons (upper). 21 rainfall in inches and the horizontal axis shows top of the figure extends along the rainfall axis with only slight temperature variations.44 35 1.00 45 7. .

and microscopic animals. animals are not directly dependent on light for their food. Experience with short wave lengths for the human skin proves that too much light may be harmful to animal tissues. 11 28 from the ' . mainthe dark-tolerant. in the light than in darkness. a photographic plate remains un- changed after hours of exposure. in the synthesis of certain vitamins These radiations are also important such as the antirhachitic vitamin D. spend their entire in the sea life in darkness. Frog's eggs do not develop normally if sunlight is excluded. 40 ness. Many. This is by no means the general rule. tained for years in their underground stalls. such as cave dwellers or animals of the oceanic depths. Arthropods and mollnsks. most favorable conditions. as well as simpler forms of life. but they grow more slowly. are killed by the . The death of desert reptiles following exposure to the overhead sun appears to be the result of overheating from radiant solar heat and not a direct effect of light. which usually live in sunlight. are found among Horses in mines. at about 2900 A. Mytilus grows more rapidly in dark- and light is definitely more injurious to marine plankton from 18 Absence of light considerable depths than to that from the surface. 3 Knowledge in this field is increasing rapidly. fungal spores. show that even the highest forms. This seems to be the meaning of the dark coloration of animals of high mountains and at times in deserts where there is little protection sun's rays.. Pigment layers at the surface of the skin afford effective protection against light. even under ^/iolet. slows up the development of insect larvae that normally live in light. Gilbert 14 has shown that the green algae found within the egg envelopes of the eggs of certain salamanders appear to have a fixed and perhaps symbiotic relation with the eggs with at least some indication that the presence of the algae accelerates Salmon eggs hatch more the rate of development of the embryos. fishes and amphibians. 17 and salmon fry are more ) active. and the presence of light retards the development of insects nor6 mally living in darkness. photonegative animals. comes occurs nearer the visible violet. . given sufficient exposure. can dispense with light. usually the cutoff many Viruses.Ultra-violet radiation reaches the earth's surface in the shorter wave lengths extending about 1000 Angstrom units (A) beyond visible The final atmospheric cutoff for the earth's surface. At a depth no greater than 1700 in.ultra-violet radiation of sunlight. quickly.22 Conditions of Existence In contrast with green plants. bacteria.

usually an inverse between the amount of oxygen and of carbon dioxide present. or nearly so. In fumaroles. seas. the death valley on the Such places are closed to animal The bodies of small birds and mammals (finches and mice) that life. The ability to live without free oxygen. of nature there given off is rarely too much oxygen for animal life. these include all air-breathers.a rich food supply can live without it. in is summer. it may displace the air near the ground on account of its greater density. often used rivers cc. there relation (pH) is of the water.4 below large cities. 30 an instance. have a large oxygen requirement with Most animals requirements. the oxygen up in waters contain 7. but that by aquatic plants has been found to be toxic to certain 2" insect larvae. In the Though oxygen is depths of the Black Sea and in 1 many Norwegian fiords that are closed by a bar. The Thames. therefore. The ability of fishes to utilize . has only 0. is is Dieng another.25 cc. fly Jarvae^ and internal parasites are good examples. where carbon dioxide escapes from the earth. Fishes avoid waters with a high carbon dioxide tension more actively than they do water deficient in oxygen. thi 1 Grotto del Cane at Pozzuoli plateau in Java. certain annelid worms. The amount of carbon dioxide present in water is apparently an important factor in the distribution of aquatic animals and is associated with the hydrogen-ion concentration In the sea and in the deeper lake waters. in a few aquatic situations. although a few animals that are able to supply themselves with energy by the decomposition of . whose oxygen per liter above London. In a narrow range of variation. nearly everywhere available.Oxygen and Carbon Dioxide Oxygen and carbon dioxide Oxygen is of primary importance 23 to life. Only a few animals are able to persist forms that can reduce their oxygen these are in such situations. where poverty in this gas is the rule. Many fresh-water lakes arc without oxygen in their depths On account of the putrefaction in sewage. scarcely affects the problems of animal distribution except among parasites and in stagnant or polluted waters. have wandered into the carbon dioxide atmosphere are frequently found at the Mofetten on the eastern shore of Lake Laach near Coblenz. Oxygen is absent. an important fact for the explanation of the differences between water. per liter below the city. the vertical circulation is very weak.and air-breathing animals. the evolution of hydrogen sulphide has combined In the intermediate depths of tropical all the available oxygen. there is a much more abundant supply in air than in water.

known to have limitations in their vertical ranges. without injury. causing gas-bubble disease determined in part 37 by the prevailing pressures. sur- empty may be subjected to a pressure of 100 atmospheres without harm. - 41 42 Food Finally. 20 In years Channel there is when mice are abundant. (See also bog waters. usually the deciding factor for density of life. turnips. For aquatic animals the pressure increases one atmosphere for every 10 m. Chapter 19. as they cannot find prey in sufficient quantity. for the number of individuals in a given area. and other vegetables in New Zealand was followed by a disconcerting increase in the native insects. if pressure suddenly removed. Larger predatory animals are unable to live on small islands. cauliflower. animals are not particular about their food and have a large menu. and perhaps at 1000. 13). atmospheres. buzzards. a sufficient amount of organic food is an indispensable condition for the habitability of an area for animals. gas accumulates in and death. Chapter 20. Then.) - Pressure in atmospheric pressure is not an imfactor environmental except in mountainous regions. 11 The planting of kale.. they are euryphagous or omnivorous. face fishes with the air bladder If the air bladder is full. . and soil acidity. The useful contrast between the terms carnivorous and herbivorous may be lost in the vari- Many ous transitional stages leading to the completely omnivorous habit. but in general we may safely conclude that the importance of this factor (pll) acting alone has been over2 3 emphasized. and there portant are associated rather with oxygen deficiency the effects of high altitudes For land animals variation than with decreased total pressure.24 Conditions of Existence oxygen when present in small amounts is decreased with the higher hydrogen-ion concentrations. i. owls. and these are the blood. Deep-sea fishes without air bladders may live normally under a pressure of more than Marine and lake fishes are 800. Even so. depth. In years when there is a richer development of zooplankton in the Engis The amount of available food lish a larger mackerel catch. gases enter the blood under the higher is pressures. This means that under these conditions they can range through the upper 1000 in. and other predators gather in the mouse-infested areas (see Fig.e.

in the earth. as in the ant. are eaten by animals. a few snails and insects feed mammals may be The upon 21 them. Stenophagy tends to limit distribution. caterpillar it if its were not so at strictly limited to a single was unknown :{f> the snail Helix aspcrsa is euryphagous man over the greater part of the earth. while the omnivorous raven ranges almost from pole to equator. when it appeared almost at once. The bird called the nutcracker is limited in Siberia by the occurrence of the nut pine. in water as well as on land. the larva of the zebra swallowtail butterfly. euryphagy to extend it. If sufficient . The distribution of the green sea urchin coincides with that of the hydroids that constitute its prinThe euphorbia sphinx would have a much wider discipal food. which feeds only on plants of the genus Sedum.. it has been able to accompany distribution of The wide most predatory attributed to their euryphagy. however.and termite-eating Myrmccophaga and Manis. of plant origin) reaches these situations. in their distribution to the areas Animals are not restricted green plants are found. they where may It is sufficient if organic matter reaches their thus occur in caves.e. there may be euryphagous and stenophagous carnivores and euryphagous and stenophagous herbivores. the lion that eats a calf equally with the fly that sucks the calf's blood. i. Most plants. which feeds only upon pawpaw. stenophagous forms are represented by the caterpillars of many Lepidoptera.Food Crows and bears 25 are omnivorous. in ground waters. Islands with a flora primarily of ferns accordingly have a strikinglypoor fauna. The flesh-eaters are thus indirectly dependent upon the plant world. such as the Apollo butterfly Parnassim apollo. some- times sharply to the species of a single genus or other higher cate7 Euryphagy is general among carnivores. habitat. tribution of plants.eating types tend toward stenophagy. The migratory grasshoppers and some caterpillars are euryphagous herbivores. Both carnivory and herbivory may be narrowly specialized or have a wide range. organic matter (always. animals may be abundant in them. mosses and ferns are little eaten. and the oleander sphinx Sphinx nerii. and in the depths of the sea. Other insects are limited to related host species such as potato and jimson weed. in genus Gottingen until Euphorbia was planted Because the Botanical Garden. Some plants are protected against being consumed by animals. and a few birds may be driven to eat them in times of famine. and they form the foundation of the food supply of animals. in the last analysis. but various insectgory.

on the contrary. the selection of animals in a given environdetermined by the habitat factor that most approaches the The more is closely even a single factor approaches a limit- ing value. designated as the ecological valence of the animal. Light and oxygen are wanting at relatively few places. favorable temperature. The amplitude of the range of the conditions of life. In Habitat-limited a few extreme cases animals may be ubiquitous. the species can live in various habitats. ment is minimum. which is stenothermal and stenophagous. and food and water supply vary in a much greater degree. or the tiger. but temperature. ecological valence. In contrast with these the oleander sphinx. is limited in habitat. quirements Artcmia in strongly salt inland waters. which a euryphagous carnivore. and these factors accordingly are the most important causes of the variation in animal distribution. The law of the minimum The basic habitat requirements together condition the distribution of animals. the fewer the number of species in the situation in ques- There may be sufficient oxygen. tion. may be conditioning factors. are specialists and do not appear in many catula. This is Liebig's "Law of the Minimum. large. When the limits for the greatest possible number of single factors are widely separated. Liebig found that in the growth of plants the food element least plentiful in proportion to the plants' needs limits their growth.26 Ecological valence Conditions of Existence Animal life is not tied hard and fast to unalterable values of the Each factor has a specific range. lying between an upper and a lower limiting value. The former have a The snail. Galba tmn- both eurythermal and euryhaline. within which an animal is able to exist. the latter a small. but the high salt content permits the . when close together. humidity. it will be limited to one or a few types of environment." This rule may be extended in a similar sense to cover the effect of the environment upon animals since that factor which a species has the narrowest range of adaptability limits for its existence. In other words. Species that can adapt themselves to varied environments can naturally be widely distributed. but the deciding factors are those most subject to variation. is which is eurythermal and It is entirely compatible with this fact that such occur everywhere on the earth where their habitat respecies may like the salt-resistant crustaceans of the genus are found. environments. are such adaptable forms. and abundant food in a salt pond. species.

"Studies on the fauna of some thermal springs in the Dutch East Indies. Gratz: 303vital Habcock. "Studies in marine ecology: III. P. Acad. 5:51-67. It supports no commercial fisheries although it lies in the latitude of Iceland. BIBLIOGRAPHY 1." Proc.. Hudson Bay furnishes a striking example of temperature as a limiting factor. Adolf. 263 figs. Philadelphia. obtained three 10 with an expenditure of 20. "Cber die Beziehung zwischcn Fortpflanzung und VerVcrhandl. "Living water. Intern. M. Alfred E. Wisconsin Expt.. 118:607-611. Brues. nor the adults are injured by low temperature. Univ. 1912.. Emerson. 176. W. Appcllof. Porfirii I. even in summer. For example. F. Berger. A fishing expedition in the eighteenth century...8 C. "Uber die Widcrstandsfaliigkeit der Tenebriolarven vom 8. The continis There ued presence of an animal in an environment depends on that developmental stage in which it has the least adaptability. Research Butt. gegen Austrocknung. 3. an extension of the law of the minimum. T. P. E. 4. Experimentcllc cntomologische Studien 7. Rcpt. Bruno. London. 73:71-95. Orlando Park.. 1907. All the waters of the bay except a shallow surface layer have a temperature below 1. A. The principles of animal ecology. Schmidt." Arch. 10 figs. Bachmetjew. Rev. which is not the lobster Homarus does not reached by the arctic waters. 1901-1907. Leipzig. "Animal life in hot springs. Biol. C. . Animal life in deserts: a study of the fauna in relation to environment. may have a superabundance of food and 27 A polluted body of water the low oxygen content limits optimum temperature. 1939. fishes with experienced fishermen from Spitzbergen." 22:87-181. Arnold: x\.. which has notable ones.Bibliography existence of only a few euryhaline animals. 44:205-253. breitung mariner Tierformen. \V. Some physical factors Biol Bull." Alice. phenomena. Physiol. but the animals to a few forms such as some Protozoa and the oligochaete Tubifex. Idem. its production and role in Arm. and K. S. 1923. Am. Zoo/. 43 figs. 1949. C.000! another application of ecological valence to the distribution of animals.. the segmentation 4 stages.." Quart. xii. C. "Metabolic water.. 5.. 1923. 1912. Sta. Arts Sri'. ges. 9. 1927. pass the latitude of the Lofoten Islands on the Norwegian coast because the postembryonic development of the larvae demands an average temperature of 15-16. Congr. Biol. Saunders: 837." 311. phyfiikalisch-chemischcn Standpttnkt aus. Rev. Buxton. Engelmann: 2 vols. 2. 6. 2:181-203." Quart. 29. Alice. neither the eggs. A. 1930.. related to the distribution of littoral invertebrates. Thomas Park.

1943. 86:327374. "The alga-egg relationship in Ambystoma maculatum. and extinction of the dinosaurs. Berlin. 1930. and 22. August. A." Econ. "The toleration of solar heat in desert reptiles. illus." Bull. illus. Rev.. The effect of sunlight. 17:101-145. 23. Erdgeschichte. Longmans: 242. evolution.s. Holzel: 2 vols. Bogert." Ann." Am. The fats. Leathes. Huntsman. 25. G. W. 25). "A new melanic lizard from Trans Jordan ia. J. the hydrosphere. "Sur la circulation moleculaire du sang chez les animaux aquati<ines. p.und Kerngrdsse und Bedentung fiir die geschlechtliche Differenzierung und die Tcilung der Zelle. Assoc. 108-119.. pi. 17. Raper. "Geography and zoology. "Fisheries research in Canada. Matheson. 1-14. Physiol. Aristides.28 10." Ecology* 25:366-369.. 54). F. 1908. 1. Sect. 11. 17:5fr-66. (3) 15. Idem. 36-41. and H. E. "The effect of light on the growth of the mussel. Parker. Borntraeger: 175. 42:31-51. 21. B. Nitsche. in 29. Perry Bull. Crozier. 1885-1895. II. John. 1 fig. R. 1936. G. Clark.. Roy. Museum Natural Hist. figs. "The vital limit of exsiccation of certain animals. EntomoL.. 26. "The utilization of aquatic plants as aids in mosquito control. Suess. Walter." 14. 98:117-123. 1:358-389. Mosauer. 16. Trans. W.. 20:709-737. vii." Science. Daniel. and some speculations concerning melanism. the critical thermal increment for the locomotion 12. "The effects of temperature on the development of an insect (Popillia japonica Newman). and James Drummond. 5: 23-28. 1903. fig. 19. Soc. Idem. Naturalist.... 9. Neumayr. Richard. 1935:137-142. ix. (n. "Ueber Correlation von Zell. 15.. 1915. 1921." ed. Whitcomhe & 9 Tombs: 375. (vol. 1-3. J. W. 64:56-86. J. Gen. and F. Am.. . et des tissus W. 11. 1. Leipzig. A. Michelberger. 16:163-190. 1944. 1924. (vol. Vienna." Proc. illus. Melchior. 18. ihre Hertwig.. 28. 1946. 1:10-17. the influence of temperature on the rate of em- bryonic development. a case of symbiosis. Ges. (p." Physiol Zool. 1904. London. Kanitz. 1940. pis. F." 24. Judeich. Lehrbuch der mittclcuropiiischen Forstinsektenkunde (mit einem Anhange: die forstschadlichen Wirbclthiere). B." 13. Cowles. "On S. Am. Fredericq." Arch. E. 11 figs.. Robert. Conditions of Existence A. E.. 23:49-62. Murray.. 33:141-143. 1914. "On of a diplopod. 27. ipi. Biol. "The distribution of organisms Intern. "Temperature tolerances in the American alligator. 31. Physio!." Ecology. Hutton. and their hearing on the habits. H. 1925. Ludwig. London. Leon. 30." Proc. Hydrobiol Hydrog. 1927. figs. "Limiting factors for marine animals: 1. F. 1920. Z. 1-4. M.. 3rd Bibliographisches Institut: 2 vols. 1905. 1922. 7:123-136. allgem. biol. 4). 1924. Canada. Zool Soc." Biol Zcntr. "Some entomological observations in California. Hall. ) 2:81-88. J. and C. Krogh. Gilbert. 20." Contribs. Temperatur und Lcbensvorgtinge. 1928. London.. Colbert. Canadian Biol and Fisheries. 2nd ed. Geogr. Animals of New Zealand: an account of the colony s air-breathing vertebrates. 1935. text figs. J. H.

(N. Idem." Syst. J. 1910. V. Plankton-Kunde." Wiss. Zool. Idem. (N. 1 map. II.. 1928. Organ. . text figs. 365 41. Helg. Geog. 35." Biol Bull. 1087. 38. Siuierlandes. Schmidt. Adolf. 9 diagr. Laboratory and field ecology: the responses of animals as indicators of correct working methods. "Ueber den Einfluss der Temperatur auf die Entwicklung von Fisch-Eiern. Peter. 265 figs. naturforsch. 5:xiii.. "Industrial fatigue in relation Physiol Revs. Wichmann." Bull. "Der Bergbach des biologische Untersuchungen.." 4. Faunistish- Int. to Heft 2.. 608. V...) 13:157-179. 1913.. 1942. "Anabiosis of the earthworm. August. "Untersuchungsfahrt des Reichsforschungsdarnpfers das Barentsmeer im Juni und Juli. Kiel. tions. Hydrobiol. Shelf ord. Sta.. Prentice-Hall: x. Gcs. Biol. Baltimore. Sitzbcr. Leipzig.." Puget Sound Biol. Reibisch. Jahrb.. "The reactions of certain animals to gradients of evaporating power of air. "Untersuchungen iiber die Fauna der Hohlen. Soc. Zool. 368. Sverdrup. 1913.Bibliography 32. 8:130-150. Pub. 1929. H.. Meeresunters." Arch. Seitz. E. 5:281-343." Wiss. "Allgemeine Biologic der Schmetterlinge. 306 figs. and Richard H. Poseidon in Ludwig. Peters. ges. Hydrog. 1 map. 1890. 1913. Vernon. 27:57-73. illus. 1905.." /. 44. 1:89-107. 43. 1912. Fleming. Y. 1926.) 6:215-231. 1912... 25:79-120. M. their physics. "Effect of light on development of young salmon.F. 40. 7 charts. E. 33." Freunde Berlin. atmospheric condi- 45. 42. 1918. "Der Grad der Beschleunigung tierischer Entwicklung durch erhohte Tempera tur. 34. 723. 12). Thienemann. Adalbert. Enttvicklungsmech. Die Hydroiden. 20:130-154. 1 pi. Exptl. 1924:113-142. "Animal communities in temperate America as illustrated in the Chicago region: a study in animal ecology.. Teubner: xv. The oceans. Smith. U. 11. Chicago. Stciicr. E. SluMiring. Martin W. Rcc. 39. N. 1:1-125 (p..F. and general biology. 37. 1915. 1922. Meeresunters." No. 36. Williams & Wilkins: xii. chemistry. 29 Karl. Johnson. SuppL.

. disadvantageous. Many flourish less groups of animals are therefore entirely absent. There the is All the phyla of the animal kingdom are represented.3. Wherever the environmental conditions deviate from such an optidifferentiation. become tions. conditions unfavorable requiring special adaptations are relatively few. and rank growth with a high tendency to variation combined else. The most favorable environment of all is afforded by many littoral areas of tropsalt content is subject to little variation. in an isotonic environment. The fauna of these seas is richer in variety of form and color than any- ical seas. with an annual variation of perature nearly and abundant food 2. Many types of organization and many individual species are unable to withstand such deviations or to transform themselves in adaptation to them. Under the stress of conditions that urgently of species diminishes. and are nourished adequately and without effort. life mum. where with the severe struggle for existence between different animals has produced the greatest amount of physiochernical conditions for animal afforded to the embryos of birds and These optimum be compared with those may mammals. which develop in the brooded egg or in the uterus at a constant and most favorable* temperature. in general. 30 demand adaptation. and approach any limiting condian impoverishment takes place in the number of major groups represented and the general diversity of the fauna. the temconstant at about 25.The Effect of Environmental Selection on Animal Distribution THE FACTORS THAT CONDITION ANIMAL EXISTENCE ARE FAVORABLE IN varying degrees at different places on the earth's surface. and others and tend less toward variation and speciation. the number Adaptations to similar environmental factors will not. only supply is brought in from the land.

6 (.3 Number of Species 34 10 1JUM1. The is families of ascidians 10 species in the tropics.4 11. number of types that can adapt themselves to the new conditions is relatively small. attests the effectiveness of The the selective process with respect to the higher categories.!)' endemic " tt 103 of which (>1 (59 l /c ) endemic " Subarctic GO" 71 " l(MHU>' r f ) 434" 635 " " - " " " " " Tropic Sulmntarctic Antarctic ei (49. species all attain their greatest number of the distribution of their 109 genera and of the shown in the following table: Species t ) (Client Arctic IJ4 of which " 2 (5. even among those of diverse ancestry. for example.) 35* (81.1- 8 .5 -20.Aquatic Life be able to take tain similarity 31 will thus traits in many different paths.) 3 (13. the lowering of the temperature results in a darkened coloration in many forms.O /C ) ( u The small number of endemic genera in the Arctic and Antarctic. Environmental selection of aquatic life Changes in marine faunae are principally associated with the decrease in temperature with approach toward the poles and with the factors accompanying an increase in depth. with a large number of endemic species. of reef corals occur in the Hawaiian Islands. the arctic plankton 1 Whereas ninety species exhibits a certain constancy of character. and the relation between the type of habitat and the appearance of its fauna becomes more obvious. In both cases the reducIn contrast with the great tion in variety of animal life is notable. commonly without pigment. and with a resulting pale coloration.4^) 503 (88. A disadvantageous environment forces common traits upon the inhabitants. variety of types of the plankton of warm waters. Such common characteristics may result from the direct influence of external factors. The Atlantic copepods in 1900 were distributed as follows: 2 Between Temperature Limits of 47.G' r ) " - *10 4* " " (7!).(> . They produce a cer- and will introduce convergent the inhabitants of a given environment.()<<) (SG. 20 only ten species of Madreporaria are found in the Bermudas.3*. the northern limit of their range. the absence of light for cave dwellers results in a fauna of creatures without eyes. c ) 173 3<> 41)" *> u 7(14.

In the salt pits at concentrated basins. gr. Bourg water to the pits contain 2 nemertines. Artemia salina. finally. and a still greater number have the majority of their representatives there. in the next series of evaporation basins ( 1718 Be) only the turbellarian and one annelid remain. If this variation is large. de Batz that conduct the salt the channels (Loire Inferieur). The number of species in the fauna is regularly de- creased with increasing concentration by the continued selection of 7 euryhaline forms. evaporated in successive shallow for the production of salt. Even Artemia. as in Cardiurn edule. 6 annelids. 1 crab. This form of selection is nowhere better illustrated ity to which sea water. only a few families are absent in the tropics. 7 snails. Similar phenomena may and Nereis diversicolor. the second series of tanks (7-8 Baume sp. the first evaporation than in salt pits. 6 annelids. in is basins contain 3 turbellarians. in total The reduction number of species with 16 depth is shown as follows in the Challenger collection: From Depths Meters 0183 1815 of: 9151849 18*9- 47443<>58 30584573 340 Below 4573 435 015 4744 000 \urnberofspeeiesofanimiils 4400 4050 710 500 similar impoverishment is caused by any sort of deviation from Variation in the salt content of sea water has an especially important influence on marine animals. A the optimum. which is especially adapted to waters of high salinity. 9 snails. great demands are made on the adaptability of the organisms concerned. becomes stunted or defective as the salt content approaches limiting values. 3 lamellibranchs. and 1 amphipod. and 7 amphipods and isopods. Macrostoma hystrix. The complete elimination of a species is frequently preceded by the production of stunted forms. . 4 lamellibranchs. and 9 amphipods and isopods. They must be protected from osmotic changes in the salinity of their body fluids.32 Effect of Environmental Selection the marine fishes of Among shallow coastal waters. 2 annelids. 1 shrimp. almost as many as are present in all the seas and rivers of Europe together. while a great number of families are confined to the tropics or subtropics. Seven hundred and eighty species of fishes are reported from the coasts of Atnboyna in the Moluccas. in the salt beds (up to 27 Be) there is only the salt-tolerant crustacean. 1 isopod. 1 crab. 1 crab. ) contain 1 turbellarian. either by an impenetrable surface or by the abil- eliminate excessive amounts of salt or water through their excretory organs.

. arachnids.. km. The reduction in number of species in such basins as the Baltic still greater impoverishment of the fauna of fresh Leaving out of account the secondarily aquatic forms in fresh water.3% in the Gulf of Bothnia (cf. draining 1. The Nile may be compared with the Obi. The Indus with 113 species compares with the Saskatchewan with 22. with approximately equal basins. rotifers. while the Mackenzie. The salinity decreases from 3% at the Kattegat to 0. a small family of sponges. iron oxide. The cheppia reaches 45 cm. ) number of species. draining 1.. has 170 species of fishes. few turbellarians. Even the fresh-water fishes..8% in the Belt. Hubbs. the optimum temperatures of the tropics have the same effect as in the seas. protozoans (especially ciliates). both with drainage areas of about 3. however. Using Giinther's figures.000 sq. fishes in lit.000 sq. are far behind the coastal marine corresponds to a waters. but otherwise marine. map at the end of Chapter 15).4 and 0.750. the former with 101 species.000. etc. all the insects. crustaceans. The two forms in the Tessin Lakes are known to the fishermen as the cheppia and the agon. there is a vanishing remnant of originally A few coelenterates. and a relatively small number of genera of clams and prosobranchiate snails enter fresh water. km. and sinks to 0. the agon usually only 25 cm. the Ganges with the Brahmaputra.000 fresh-water species. hydro- gen sulphide. Additions to the water such as carbon dioxide. The size of shad of the lakes of north Italy shows that fresh water less is favorable than that of the ocean. The agon (Alosa finta var. require special adaptations and thereby . in reckons about 30. phenomena appear with decrease of the salinity of sea The Baltic offers an excellent example. the latter with 45. annelids. The cheppia (Alosa finta) is a migratory fish going up to the lakes to spawn. and pulmonate snails. and nearly all the annelids are more or less altered forms of North Atlantic species. and gastrotrichs have found the fresh-water environment favorable. change transforming upon the marine fishes. A modern estimate (Carl L. which have entered it from the land.500. and bryozoans. s Increase of temperature above the optimum impoverishes the freshwater fauna in the same way as does decrease.000 sq.e. as in marshes and in regions without outlet to the sea.000 marine and 10. a aquatic forms. Fresh.Aquatic Life be observed wherever the salinity of waters increases. G In fresh waters. lacustris) has become permanently resident in the lakes. km.water fishes. mollusks.. This a All has selective the and effect fauna. humus acids.. has about 23. i. Parallel 33 salt water.

It is the general rule for air-breathers that their development favored by sufficient humidity. birds.34 react selectively Effect of Environmental Selection upon the fauna. are inhabited by a fauna dwarfed by the severe habitat conditions. nemertians. such as Gastcrosand teus. tiny crustaceans. and these alone compose three-fourths of the known species of animals (about a million de1<5 scribed forms taceans. and worms are to be found. raise the number of land animals probably to at least four-fifths of the total. phyla is absent in the ocean ( even if the Gastrotricha are regarded as of this rank). terrestrial animal life exhibits an impoverishment Not one of the animal in wealth and variety of structural types. Of the 186 species of rotifers in Galicia. land snails. The myriapods. rich in humus acids. and many subphyla are still greater number of entirely absent among the air-breathers. sponges. Lakes very rich in iron. whereas the only prominent classes entirely absent A in the sea are the Onychophora (Peripatus). reduces the number of species.000. 11 The number of species of animals decreases . live in bog waters. 17 Environmental selection of land animals Terrestrial animal life life. as in Ritom Lake. bryozoans. Tinea. arachnids. myriapods. The carp find these waters less favorable. and the and mammals. Cobitis. The number of terrestrial species greatly exceeds those living in water. Admixture of hydrogen sulphide. coelenterates. 12 Greenland (half as large) has only 599. constantly demand adaptation. relatively constant temperature. land crusin addition. in New Caledonia. on the other hand. The number of species of insects in India is 40. Canton Tessin. but in types of organization. and amis phibians. while protozoans. toward the poles. flourish in such water. These conditions are found in optimum combination in the openings in tropical forests like those of the Amazon. a small fish of the genus Gdlaxias. Decrease of moisture and temperature. animal life is least abundant on land where the habitat conditions approach the limiting values. or of New Guinea. ctenophores. as in high mountains. Rotifers. in steppes. reptiles. and especially great fluctuations of these conditions. Only a few fishes. the Congo. classes are absent. ) . small snails. not in number The great majority of insects are terrestrial. is also impoverished as compared with marine of species or genera and certainly not in the numbers of individuals of special forms. 100 may be found in bog waters. In spite of this fact. and abundance of light and food. or in deserts. As in aquatic habitats. and pike and trout avoid them. and echinoderms.

They live by eating flesh or fungi. The uniformity of conditions presented by the African grasslands south of the Sahara results in extraordinarily wide ranges of various species of mammals. Chapter 20) exhibit the opposite relation. Caves. or reptiles. poor in species and individuals. is much more variable than the oceanic herring that keeps to the open sea except for spawning. The number of species. The size of species that range from the lowlands to the tops of mountains often decreases with altitude. which are reduced in size. physical environment effective. myriapods. i. and amphibians. and vertebrates. often with limited ranges. whether 9 amphibians.. notably exemplified in the plethodont salamanders of the Appalachian Mountains. all have representatives in the sea. or matter originating outside the caves. this area may be stated as "Senegal to East Africa and the Cape of Good Hope. on account of their relatively low temperature and the absence of light arid plant food. seems to depend on the variety of habitat conditions and on the adaptations required by them and on opportunities for isolation. upon the transformation of species is extremely factors do not seem to have been the cause of Physical the development of the primary divisions of the animal kingdom. support only a restricted fauna. exthe rain-forest. In the tropics and subtropics the average size of cold-blooded terrestrial animals is cold zones. reptiles. Warm-blooded animals of larger than in the temperate and (cf. birds. the inhabitants of unfavorable environments are so changed by their physiological and structural adaptations that they are recognized as new species or even The indirect selective influence of the as new genera and families. the mollusks. or changes typic varieties. arachnids. into dwarfed forms or phenoFor the most part. with wooded mountain slopes clusive* of 1 . however. in the lighted zone of the sea. with a north-south axial mountain range 4000 meters in height. which is subjected to diverse habitat conditions.e. them only to a slight degree. The three principal groups of ter- restrial animals. arthropods. Varied mountainous topography tends to the development of a more varied fauna. which apparently developed in the same rather uniform environment. This size relation applies to ancient as well as to insects. however." 1S Wide ranges throughout the Amazonian and African rain-forests are also the rule. The herring of the coast. In some places the selective action of the environment leaves species with special powers of adaptation unchanged in their new habitat. snails. modern groups. Formosa.Land Animals 35 steadily with increasing altitude on the mountains.

-.. and he who escapes Charybdis falls into the Where the struggle with the physical forces of nature becomes more This is severe. Even predatory animals are compelled to protect themselves against much more enemies of jaws of Scylla. which are the more striking the more closely the environmental characters approach limiting values. in the steppes. Effect of Environmental Selection coastal plains. some of which are com- pletely restricted to fresh waters. where terrestrial animals of temperate zones. and in the polar regions. in deserts. petitors is many kinds. like the sturgeons. the strugfor existence between the different species of animals will be much gle more violent than in areas where the fauna is poorer in representatives ment. is remarkable for the range of its climatic conditions and for a fauna rich in number of species. true in fresh water. as a result of the reduction in number of types. and the opportunities to get the better of a competitor are much more numerous. optimum less and less important and are restricted to special habitat conditions.it has almost as many species of birds (128) and and mammals (35) as Japan (165 birds and 40 mammals). Accordingly. allowing only more or less similar forms to Thus arise the common characters of animals living under pass. conditions common to a fauna are like the floating arrangements of plankton or the adhesive apparatus of the inhabitants of surf-beaten rocks. in the polar Such common traits regions. So do the ganoids. while others. similar conditions.g. are difficult to discover in animals of large bodies of water or in the In the tropical seas. in the tropics) there severe competition for the same goal. most notable in deserts. Among the bony fishes the more primitive soft-rayed forms. and in inhabitants of temporary pools. is In the former case (e. in competition with the spiny-rayed Acanthopter- . According to Wallace. conditions such adaptations to the environment are reign. Many animals are able to maintain themselves in physically unfavorable environments after they have given way elsewhere to more modern competitors. Another interesting mutual relation may be explained by means of the factors that condition the wealth of form in a given environ- Where the number of morphologic types is greater. Thus the last representatives of the lungfishes persist in fresh water. Adaptive selection The necessity for definite adaptations acts upon the fauna like a sieve of definite mesh. of diverse structure. repair to them to spawn. the struggle for existence between animal comreduced.36 and ravines.

adaptation to an environment enables an exceptional. in regions equally well supplied which one has other environmental conditions favorable and the other unfavorable. rotifers. in both glacial streams and hot springs. plowed. 120 different to species. 14 An intensively and scientifically managed fish pond. which night in Borneo. which is annually drained. the number of competitors for the food supply will be reduced in the new environment. as in the North. is more than compensated by the enormous number of indispecies viduals of the few flagellates. During one ijl collected 158 specimens of moths. is unproductive for a collector. animal to flourish under changed conditions that are unfavorable for unadapted animals. and when a sufficient food supply is available the numbers of become enormous. sists and Galba truncatula occurs not only in very saline waters but also in almost fresh waters. cladocerans. In the tropics the favorable conditions (combined with the intensity of the competitive struggle) permit almost unlimited speciation. Thus of However individuals of the few adapted species may with food. Koningsberger belonged the individuals of many . often of reduced size. in the oceanic depths. The brackishwater fauna is characterized by the presence of few species. temperatures are necessary for steno thermal cold-tolerant animals like Planaria alpina or the trout. Low There are also broad adaptations resulting in increased bodily reNereis diversicolor persistance to adverse environmental factors. where four-fifths of the physostome species are found. Wallace 13 states that in Java. this crustacean is often the only animal of considerable size. and freed of large plants. as in Nereis diversicolor in saline springs or in some protozoans in hot springs. and in fresh water. the number of individuals of the species will be in inverse proportion to the number of the species present. The number of individuals in may be so great that the water is a thick broth of Artemia. conditions Although the persistence of some species under increasingly adverse may produce degeneration. but with enormous numbers of individuals. and cyclops that do persist. but such waters species may not be abundant. manured.Numbers of Species and of Individuals 37 ygii. the unproductiveness in invertebrate forms. have maintained themselves everywhere under more adverse environmental conditions. In salt waters inhabited by Artcmia. of species Numbers and of individuals the adaptation to adverse conditions may have been reached. perhaps accompanied by the larvae of the brine fly Ephydra. these cases are In general. for whom a neglected pond is a rich source of supply of For the fish culturist.

Amblyrhynchus Photo by R. The 311 Fie. Herbivorous reptiles are not numerous. but the numbers of individuals of each species are extraordinarily countless great . numbers of bison that dominated the plains of North America be contrasted with the great numbers of species of antelopes may on the savannas of South Africa. where they do not suffer from mammalian competition and where the total number of vertebrate species is low. but of the Galapagos Islands.v. men of many numbers and secured only one specinumber of Bornean bird species is which is 13 times as large.{ never saw the well-known birds of Borneo in species. of Narborough Aggregation of marine iguanas.38 by Effect of Environmental Selection collecting on the flowers and shrubs of cultivated districts. though the almost equal to that of Europe. 2. when we remember Amblyrhynchus ( thousands of large tortoises terrestrial the well-beaten paths made by the the many turtles the great warrens of and the groups of the ) marine species basking on the coast-rocks of every island we must admit that there is no other quarter of the world where this order = Conolophus . Whitehead 2. each of which is represented by smaller numbers of individuals. Island. Darwin wrote: "The species are not numerous. II. the California rm/fi/M. Galapagos Islands. on tin* coast Beck. 100 species of spiders with one specimen each are more easily secured than 100 specimens of a single species. . Courtesy of Academy of Sciences. .

A. 1949. 2 maps (1st ed. Victor. Introduction to the study of fishes. London. number New Zealand 11 ' is only about a dozen. The salamanders of tlie family Plethodontidac. figs. 46). Chun. physical conditions. Ferronicre. the species represented by few individuals are so numerous that one-half of his total list makes up less than a tenth of the individuals. 209. llandlirsch. 1910. Ges. "Relative abundance of some Panamanian snakes. 60:160-185. Niigcle: 1-64. 7. 5:lxxx. 6 pis.Bibliography 39 3 replaces the herbivorous mammalia in so remarkable a manner. P. Analysis of such mass collections of snakes from various parts of Panama by Dunn exhibits proportions of abundant and rare species r> not very different from those in a collection from a temperate region. 'Beagle* under the command of Captain Fitz Roy." 30:39-57.. Birds of Malaysia. (p.. Journal of researches into the natural history and geology of the countries visited during the voyage round the world of H.N. T. during a clearing operations produced no less than 407 specimens in 5 years.. 1897. 551. . map. Charles R. Stuttgart. The relations of numbers six of individuals and numbers of species Alfred in the tropics may be obscured by the availability of cover. Murray: 3fl. 11 text figs. Zoo/. Carl. It must be remem- bered that a large number of individuals is a correlative of a small of species only when the conditions of food supply are Thus the number of species of butterflies in especially favorable. but a collection from forested Darien made in the course of land- Emerson." Bull. "Geographical distribution of Atlantic copepods and their Ofv. 1 pi. observed only a single specimen of the fer-de-lance (T rimeresiir us atrox). soc. \ViVn. 1839). 321 9. Delacour. ton.. C. R. K. 1880. Mass." Verhandl. Bot. R. Dunn. Ciinther. 57:139-144. Jean. 85 maps. Smith College: viii. L. NorthampEcology. figs. 1890. (pp. de 1 la Georges. Vetensk. "Etudes biologiques sur les zones supra-Iittorales Loire Inferieure. 382. Clrve. suisse." Similar examples could be increased indefinitely. 576. 4 pis. ouest France* (2) 1:1483.. Faune vert. 1890. BIBLIOGRAPHY 1. 1 map. Edinburgh.. Akad. Black: xvi. 1926. "Einige interessante Kapitel der Paliio-Entomologic. For/i. sci. New York.. Darwin. 1901. Fatio. E. xvi. 720. Die Bezieliungen zwischcn (Jem arktischen ttnd antarktisehen Plankton. Anton. months' stay on Barro Colorado Island." 6. G. illus. 2. about one-tenth of the species make up half of the individuals... 8. 1947. 441. "Ilistoire naturelle des poissons.MS. New ed." 3. 84 4. 1900. On the other hand. Idem. 5. Macmillan: xvi. 259).. nat. with no especial abundance of specimens.

S. Sarasin. P. Indian Set. 106). John.. 3. 1-27. Zoo'L. illus. (p." Bronns Klassen u. Balu. 81-87: 1281-1488. 468 figs. 3rd Intern. 1935.40 10. Idem.. Fritz. R. Museum Natural Hist. 184 figs. R. The insects of Australia and Neiv Zealand. Leyden. 23. 96 pis. L. Pt. The Malay Archipelago: the land of the orang-utan and 19." Grjnland... Mobius. H.. Pratt. W. 28). Buitenzorg. R. John.. Philadelphia. Proc. xx. Koningsberger. 13. M. 59:ix. Macmillan: 1893. R. 1909. 1895. figs. . 1883.. K. Murray. Neu-Caledonicn Basel. Vaughan. Congr. und die Loyalty-Inseln Reise-Erinnerx. Henriksen. Suppl. including a revision and attempted solution of the problem of geological climates. 7-32. S. A. 14. Java ZoologiscJi en Biologisch. 183). 17. 1:1-663 (p. London. "Contributions to the herpetology of the Belgian Congo---. 1869. 1938. and Friedrich Heincke. 22..." Bull U.. 11. 1895. "Presidential address. Karl A." C. 12. Paray: 15.. London. 206. Medd. Wallace. 39:385-624. 427. 18. Ifg. 32 pis. Blakiston: xviii.. 3:99-117 (p. 207). 1 map Schmidt. 1919. C. 21. 974 figs. Gurney & Jackson: 317. 11-21). "A revised index of the insects of Greenland. Section of Entomology. A. Ordnungcn. Georg & Co. 2:201-246. 51). Island life. Whitehead. ungcn eincs (p. 16. 1926. 1917. 1 map (p. Natl Museum. 1907. Manual of invertebrates. Die Fische der Ostscc. or the phenomena and causes of insular faunas and floras. London.. Angus & Robertson: xvi. pis. "Tunicata (Manteltiere). "The general conditions of existence and distribution of marine organisms." Congr. 185). 44 pis. Tillyard.. Naturforsclicrs. 854. K. 1939. Sydney. iv. Effect of Environmental Selection Hartmeyer." Bull Am. 563. J. Exploration of x. J. T. 560. "Recent Madreporaria of the Hawaiian Islands and Laysan. NortJi Borneo. the bird of paradise. illus. Mount Kina 21 text figs. Macmillan: 2 vols. 119:1-111 (p. 2nd ed.. 20. 1911-1915.: 284. Hussein. (pp. Berlin..

involved. internal whose conditions of life somewhat resemble those of aquatic classification is into terrestrial animals. According to Classification of Animals Characteristics the Most General of the Environment THE OLDKST ATTEMPTS TO CLASSIFY ANIMALS. . independent of morview of the obvious structural adaptations them into air-breathing and water-breathing forms. not only the animals living on the earth's surface. include. and aquatic and not \yatcr. birds. its justification. and bats). AS IN THE BIBLICAL ACcoiint of the creation. Terrestria. and some earthworm genera. and varying humidity. All animals Another environmental whose bodies are surrounded by air. whose ancestors have always lived in water. Allurus for example. It must be admitted that there are various intermediates between these two groups. but also the subterranean and wood-boring forms.e. Aquatic animals. and Vplatilia. high oxygen content. Frogs in the northern hemisphere are air-breathers in summer but strictly aquatic during their winter hibernation. animals. An ecological classification has in phological taxonomy. are in this sense terrestrial. i. Many jiewts are aquatic in The common earthworms may be land.airjjireatliirig types. all of which are subjected to the influence of the air with its low density. Pliny in the first century A. These preserve summer and hibernation entirely 41 .D.numerpus : . in a broad sense. immersed in water in the event of continued rains. A primary division of an ecological classification of animals separates parasites. are based upon their habitat. are sharply distinguished from ^ which have re-entered the water from the land. The primarjjy^aguatic forms.4. divided animals into Aquatilia. are true aquatic animals.. and the flying groups (insects. will not be discussed here in detail.

illustrate an approach to sessile feeding habits. with which we are immediately concerned. tides. obtain their oxygen from the dissolved supply in the water. 10 m. and they are thus able to dis11 Ant-lion larvae. for all types of structure Animals with gelatinous^ bodies permeated with water. there are fewer limitations to the organization of aquatic animals as compared with Marine animals terrestrial forms.. worms (Alciopc) and their larvae. Tridacna. The giant squids attain a body length of 6 in. Heteropoda among mollusks. an extremely wide distribution of many marine species of animals. as The greater density of the &que_pus medium helps to j^oikilothermal.. for their food supply consists of suspended organisms brought to them by currents. and such forms are found among the most diverse groups: medusae. are not required at the exposed surfaces such as the skin. The unbroken connection of of the sea water by means the oceans and the continuous diffusion of currents. while the blue whale (Balaenoptera musculus) is the largest of living mammals. All are air breathing. in characteristics associated with terrestrial addition to an unmistakable record of their racial history. The primarily aquatic animals. There are no other positive characters common to all aquatic animals. non-parasitic) sessile animals are mostly confined to the aquatic habitat. Considerable variations occur only in limited This widespread uniformity of conditions is accompanied by areas. with arms of 11 m. terrestrial forms. gills. by which the concentration of the body fluids might be altered. the whale-shark. special protective structures to prevent the osmotic exchange of substances. Aside from the arrangements for breathing the oxygen dissolved in water. salpae and eel-larvae (Leptoccphalus) among the chordates. and a weight of 200 kg. is the giant among bivalves. and storms effect a general equality in the composition and amount of the substances dissolved in sea water. and mucous . ctenophores. among arthropods the Japanese crab (Kacmpfferia kacmpfferi) has a limb spread of more than 3 in. occur only in water. Mycetozoa excepted. with a greatest diameter of 2 m. Since the body fluids of marine invertebrates 13 and of sharks arc approximately isotonic with the sea water in which they live. or more in length. among pense with locomotion in search of food. represents the maximum for fishes. their bodies and thus makes it possible to dispense with or support to lighten special supporting structures. Most groups of animals reach their maximum size in the sea.e. are represented in the water...42 numerous Classification According to Environment life. Free-feeding (i.

1 a fishes. membranes In this respect fishes are independent of the sea water. swelling the body. or excretory organs that are capable of excreting the water as fast as it enters. In Anodonta these fluids contain ten times as much dissolved material as the surrounding water. covering of the surface of many aquatic forms. Invertebrates in the Mediterranean accordhave a somewhat higher degree of salinity of blood than those ingly of the Atlantic and North Sea. and they maintain osmotic balance by actively drinking sea water and eliminating excess salts. such as the This of slime from the skin of an eel causes the osmotic pressure of its blood serum to vary with changes in the osmotic pressure of the surrounding water more than it does in a normal eel. The mucous coat forms an effective barrier to prevent the exchange of water between the outer and inner media. the meaning of the gelatinous covering of buds of aquatic {7 The diatom Thahissiosira has a mucilaginous covering that plants. and interfering with the normal functioning of the protoplasm. The removal it against the varying salt content in the water. probably protects them against the entry of water. insects and arachnids with . although not as high as that of sea water. 28 Among secondarily aquatic animals. diluting the body fluids. As a matter of fact. In the sharks and their allies this osmotic balance between the blood and the surrounding medium is maintained by the admixture of a considerable amount of urea (2-3%) to the body fluids. Fresh-water animals The primarily aquatic fresh-water animals are without doubt derived from marine ancestors and must be supposed to have inherited from them a similar molecular concentration of their body fluids.Fresh-water Animal? 43 of the gut. membranes that have a capacity change in permeability. there must be devices the. In contrast. To enable fresh-water animals to exist. the bony fishes have a much lower molecular concentration in their body fluids than does sea water. for that prevent the entrance of water. A continuous stream of water must therefore diffuse through semipermeable body membranes. protects snails mucous and is In addition to such permeability changes of the epidermis. Capacity of the skin to adjust its permeability with respect to the surrounding medium is an anti-osmotic adaptation in some euryhaline animals. among such fresh-water invertebrates as the pond clam Anodonta and the crayfish Potamobius. which have a lower salt content than sea water. the body fluids have a higher osmotic pressure than that of the water in which they live.

it will continue to live. no contractile have vacuoles. When the animal is immersed in tap water.44 Classification According to Environment seals their exoskeleton. If a fresh-water amoeba is introduced into sea water by gradual increases in salinity. a paramecium own volume of water in an hour. All fresh-water Protozoa have one or more contractile vacuoles.O 0.4 exist: ls % XaCl water in in 0. :ili absorbed by a frog through its skin and that about the same amount is excreted through the kidneys in a given period. have a protection against the influx of water through the acquired during the terrestrial life of their ancestors. body surface. in by osmosis.25 0. At about 20 C. which constantly discharge water from the body. but its con44 Intractile vacuole ceases to function and ultimately disappears. an isotonic medium it is usually difficult to recognize the bladder on account of its small extent. At 20 C.. from snail blood. the following relations 0.8 1.82 1. on account of decreased competition.2 9. This shows that the water diffused into the body is being removed by the ncphriclial system.88 G In Metazoa the kidneys play the same role in removing water taken In cercaria parasitic in Galbai. and this primary condition for adaption to fresh water is not equally at the command of all marine animals. Marine which live in a medium isotonic with their parasitic protozoans. the Y-shaped bladder quickly becomes evident on account of its distention.5 18. down crease in the molecular concentration of the surrounding water slows the contraction rhythm of the vacuole and Irsscns the amount of water pumped in out. Isopods.e. Such marine forms as possess the basic requirements for adaptaonce find favorable opportunities in brackish water.00 IGiJ. No conclusive data on this subject are available for selachians or bony fishes. Relatively few tion to life in less saline water at brackish-water forms can The relatively small make the further advance into fresh number of species of primarily aquatic water.8 2.08 body volumes 4.1 Contraction period Excretion per hour seconds (>.3 24. excretes almost five times its and contractile vacuole counteracts the osmotic influx of water is furnished by experiment. It appears that the ability to increase the rate of water excretion would enable a marine animal to enter fresh water. Adaptation to rapid excretion of excess water is widespread.75 1. and whales and with their horny epidermis. and decapods of fresh water agree in having the antennal excretory organ larger Overtoil 2ft has shown that water is than in their marine relatives. fresh- . Complete proof that the body fluid. amphipods.

and Romer. The host of sponges is represented only by the small family Spongillidae. Philippines.3% salinity). groups have gained wider distributions. arachnids. otherwise the phy- worm). those secondarily aquatic. The fresh-water annelids include a few leeches. and the Syngnathidae. West Indies. well separated . but the nemertines are very few. such as Hydra and Cordijloplwra (the latter more commonly in brackish water up to 1. from paleontological considerations. in the form an important element Marine invasions of fresh water The immigration for ages and still continues. the Elopidae. Rotifers and gastrotrichs are present in much greater numbers than in the sea. but like many there are a very ties. such as insects.Marine Invasions of Fresh Water water forms shows that not tion. and Samoa) and the Palaemonidae (Europe. Of the coelenterates with their wealth of forms one encounters only a few hydroids. dominate the life of fresh water. 45 many animals are capable of such adapta- A natural capacity for adaptation to fresh water seems to exist some genera and families. fresh-water animals these tend to have a world-wide distribution. Echinoderms and tunicates are entirely absent from fresh waters. and representative members have independently accomplished the transition to fresh water in widely separated regions. the Americas. and few fresh-water medusae in widely scattered locali- The flatworms are relatively well represented. The more or less universal fresh-water forms are usually sharply defined groups. and a few quite lum is poorly represented. Bryozoa are well represented in a single family of Entoprocta. 27 of marine forms into fresh water has taken place Fresh-water animals whose close re- lationship to those in the sea indicates relatively recent entrance into Older the new habitat are naturally limited in their distribution. Despite the presence of these basically aquatic animals. " on physiological grounds. and Africa) may be mentioned. the Gobiidae. A few important families and even orders of fishes are predominantly inhabitants of fresh waters. the somewhat uniform group of limnicolous oligochaetes (aquatic relatives of the earth- isolated species of polychaetes. have concluded that fishes as a whole originated in fresh waters and later invaded 1 :i. and pulmonate snails. with rhabdocoel and triclad turbellarians. Homer Smith. primary fresh-water fauna. Among the higher crustaceans the genus Atija in (Cape Verde Islands. Examples of such groups among the bony fishes are the genus Coitus with its relatives. especially Entomostraca. Crustaceans.i the oceans.

Among fishes in the Indian to Florida. brackish. it. in water entirely fresh. Leander. crustaceans. ecologically speaking. The fresh waters about the Gulf of Bengal. Belone. The relations are still more evident among the In the islands of the Indian 4 Ocean . of 22 species of selachians occurring in fresh water. The species may occur in the ocean also. fresh-water groups are related to the inhabitants of the neighboring seas in various degrees. paludosus occurs in fresh-water streams and lakes as far west as the Chicago area. and salt water. though still subject to movement. Palaemon carcinus even in all three. water bryozoa. Of the fresh-water selachians. ascending the European streams to spawn. The marine genus of snails Tectitra has a single fresh-water species. has established itself in the Lake of Lugano.. Nassa by Canidia. and fishes. Palacmonetcs the 9 common prawn of the Atlantic coast from Massachusetts while P. the islands of the Malay Archipelago.46 Classification relatives. ' Lates calcarifer. . the freshpendent phylogenetic history. or fresh-water genera of families otherwise marine may Such relations all indicate recent immigration. in southeast Asia. Blennius. Alosa finta. Madagascar. occur in limited areas. Finally. According to Environment from their marine such groups. lives in fresh. other and marine forms occur in Trinidad the rock-boring Pholas tidal 18 km. Tcctnra flnminalis. of marine animals into fresh water continues in recent time primarily in the tropics.1 confined to Ocean. Of the marine genera Syngnathus. many decapods. the fresh-water species may belong to genera otherwise marine. Among gymnotids. The marine genus Cerithiwrt is replaced in the 7 fresh waters of India by Brotia. with a long indeare the fresh-water sponges. or S. only 7 arc 11 A species of bass. and the Ostariophysi among the bony fishes ( characins. Penaeus. Numerous examples of regional fresh-water representatives are found among mollusks. and others. in the Irrawaddy River. Where immigration is recent or still continuing. Among mussels. 14 A variety of the shad. The newer inhabitants of fresh water have not had time to diverge greatly from their marine relatives. and only 12 a few beyond 30.only 4 out of 20 genera are confined to fresh water. and tropical America no species is Pseudograpuws. single species are confined to fresh water. cyprinids. the rest live in the ocean as well. and silurids). marine genera vulgaris is * have single species in fresh water. Caridina and Palaemon live principally in fresh water but have species that occur in the sea and in brackish water. from the sea. found beyond latitude 35 N. The immigration for example.

and below greater depths a uniform salt content of 32. which have been separated from the sea by the rise of the land.. which are frequent at certain places and seasons in the tropics. from the North Sea on it occasionally ascends streams to spawn. The lower stratum is life on account of the presence of hydrogen sulphide. In 1888. from 6 to 12 m. to fresh makes the transition water less difficult. will survive and will then remain as forms with marine relations in an association of fresh-water animals. of Kildin. four or and Pycnogonum. with this <? without animal Gammarns locusta (an inhabitant of the marine beach) on the shore. there is a rapid increase in salinity. snails five tunicates. The upper stratum contains daphnias and fresh-water copepods. the middle layer contains only marine animals. 2 Lapland. 5' 25 Another process leading to the production of fresh-water forms is the freshening of arms of the sea that become cut off from the ocean. characteristic of so-called hard waters.Although now without visible oceanic connection. others. as in the lakes of Finland and south Sweden. is a lake of this type in process of formation. a sea star. this marine fauna consisted of two or three species of sponges. and about the Adriatic it has become in part permanently at home in fresh waters. Separation of arms of the sea by tongues of land is frequent along the French Mediterranean coast and in the northern Stenohaline marine animals caught in such basins and Adriatic. and other mollusks. For this reason its waters are stratiwith fied. annelids.5/1 prevails. by reducing the euryhalinity salt among marine content of the surface waters. left thereby subjected to the influence of inflowing fresh water will be destroyed. These forms are marine relicts. The small variation in temperature of tropical streams probably facilitates the entrance of marine forms. sea anemones.Marine Invasions of Fresh Water are rich in 47 new immigrants from the surrounding seas. it is still so connected at by seepage water. Ten years later a number . It is also possible that the tremendous rainstorms. a few bryozoa. codfishes. or in the lakes of the south Russian steppe. The presence of large amounts of calcium. and such Lake Mogilnoje on the island lakes are termed relict lakes. which were behind by the lowering of the surface of the former Sarmatian Sea. This may occur through changes in the strand line. however. on the Murman Coast. completely fresh water at the surface and to a depth of 6 m. help to select for animals. In Scandinavia this species spawns among reefs in the sea. Temperature may have some- thing to do with the varying behavior of Alosa finta. in association with its spawning migrations.

and Pallasiclla 34 American examples include Mysis (juadrispinosa are reported. but . the only forms with marine relatives are Mysis rclicta and Limnocalanus macrurus. skin. and mucous covering of the acters of fresh-water animals. others attain a large size of 20 to 30 Ib. The fresh water Alosa of the Italian lakes is smaller than the migratory form. although their remains were still present on the bottom. whereas in the Finnish lakes it grows only to 15 cm. emigration to the sea the fresh-water species are smaller than the marine. In the lakes of southern Sweden. 8 In the bass genus (Fig.. solar is known to have fresh-water colonies The that are merely overgrown parr. in comparison with that of the brook trout. permeability of the body membranes. Pontoporeia affinis. It has been established that the growth of the salmon. The causes size is of this phenomenon are not yet fully known. The Atlantic salmon S. are primary char- A number of additional characteristics of animals in fresh water indirect influence of this medium on have probably arisen through the direct or their organization and may be regarded as secondary. never more than a foot long. 80 Another contrasting secondary character is the decrease in the numis ber oFeggs in fresh-water animals. The land-locked salmon of Maine and New Hampshire remains smaller and stouter than the form from which it is derived. rule has exceptions.48 of the Classification According to Environment marine forms were no longer to be found alive. Thus the hydroid polyp Cordylophora lacustris is smaller and has shorter stalks in fresh water than in brackish. reduction is in connected with a departure from the optimum. only Mysis relicta. The special characters that enable animals to live in fresh water. is notably accelerated by all its Ambassis. 21 The smelt Osmerus cpcrlanus of the North and Baltic seas reaches a length of 30 cm. the lake trout from the Great Lakes and the huchen of the Danube are as large as any migratory salmonids. The nature and abundance of the food one of the principal factors responsible for these differences. but that not an explanation. yi Marine relicts in such lakes often become very few in number. This applies to nearly all forms with the exception of the rotifers. rclicta in Lake Michigan and a nereid worm in Lake Merced near San Francisco. A by yolk-masses goes hand in hand with the decrease notable enlargement of the eggs in number. 3). such as regulative activity of the excretory system. The first of these secondary features is the smaller size of freshwater animals compared with their marine allies. In the lakes of the Baltic divide.

each gonophore. tions that fresh Gobius niger. salmon trout S. Thus the flounder Pleuronectes flesus. fario. salar. 24 The oyster plantations of the river mouths cm. in salt water. for example entering the eastern Baltic from the North Sea do not spawn there. trout S. diameter. The small crustacean Palacmonctcs varians in specimens of equal size (4 cm.5-mm. does not become A number of fishes Coitus htibalis and sexually mature in them. trutta. has 321 eggs. There are indicawater checks fertility. and in total mass the fresh-water . length). which frequently ascends the European rivers. 38 The fact that egg size increases in fresh in species that also occur in brackish water is especially evident water and in the sea. The egg size is in the proportion of 1:27.5-mm. Growth curves of Atlantic salmon S. and only 25 eggs of 1. of little over 0. 100 90 80 Salmon after Migration* 70 60 50 40 30 Salmon and Sea Trout Trout after Migration 20 10 before Migration Slowly Growing Brook Trout 5% Years FIG. 3. would die out if they were not constantly The brackish-water polyp lacnstris in fresh water suffers a decrease in the number Cordylophora of gonophores and an approximate halving of the number of eggs in replenished with spawn from salt water. diameter in fresh water.Marine Invasions of Fresh Water whether this is 49 a cause or an effect is unknown. and European of the west coast of France After Duhl.

The influence of salinity on the amount of yolk simply gives a more intimate explanation of the assumption of 11 Sollas. so that the small number of eggs cannot in this case he a result of general degeneration.e. the egg size increases with reduction of salinity of the water. and a free-swimming animals young In larval stage is thus suppressed to a greater or lesser degree. In most fishes common to the North Sea and the Baltic. would always be carried to the ocean by the current of stream. and stout and 5. like those of protection against dilution . and the yolk content of the eggs of such species can have no significance for the process of acclimatization. These two explanations are not incompatible. however. larvae arc slender and Pahiemonctcs varians. and in copepods and polyphemid Cladoccra. Fresh-water animals have eggs with a dense covering. 17 Among fresh- water forms free-swimming larvae are present in the triangle mussel.901 . Another explanation for the absence of larvae in so many fresh- water animals has been advanced by Sollas. The eggs quire some of fresh-water animals. like the animals themselves. the imly animals that could persist in fresh water would be those whose eggs react to reduced salinity as do those of Palaemonetes.50 Classification According to Environment form has produced twice as much of egg material as the marine.s thai they tried to enter. the slight increase in size the fresher water suggests a mere swelling of the eggs in the weaker of yolk in fresher waters has the result that the hatch at a more advanced stage. Animals may. would be able to enter rivers from the sea. as happens with Drcisscna. 1 Kgg diameter . which is carried by ships. be transported directly into fresh water. starfish. which are capable of only weak active movements. zoe'a 4 mm. sojourn in brackish water. and this lined. Entry into fresh water is for the most part preceded by a would affect the eggs as above outAccording to this supposition. more alvanced state in brackish than in salt water. He believes that freeswimming larvae.95U mm. for example. 4 The herring in Schlcswig hatches in a noticeably long in fresh water. with large-yolked eggs and abbreviated development.ttKftV.45 1 17. the young The abundance long at hatching in the marine representatives. i. and that therefore only animals without such larvae. and swelling by the inward Marine forms may have completely naked eggs.. Dreisscna poly?norpha. in the flounder Plcuronectes platessa: Salt content 10. re- diffusion of water. Motclla cimbria. in In the Baltic salt solution.5 mm.870 .JJ 1 la.

total land surface is only 148. amounting to 330 contractions per second fishes. which even in favorable cases. Less than 7 ec. or the eggs may have a gelatinous covering. four-fifths of the known species of animals are terrestrial. as in planarians. or their eggs may be enclosed in a thick-walled case. Invertebrates of this kind are the Onychophora. On land.Terrestrial Animals 51 or the river crayfish. and land planarians. have led to the establishment of new and successful groups of ter- Many A restrial came animals. deep. is unknown among is although extraordinary muscular efficiency water-breathers.. The oceans. Among vertebrates a number of fishes from various families and orders il- . Attempts to enter the terrestrial habitat that have met with a limited or partial success are usually confined to iso- lated genera or to small groups.892. Terrestrial animals accordingly live much more intensively than the primary aquatic forms. Hydra Terrestrial animals The Terrestrial animals contrast with the aquatic forms in many ways. land leeches. but for the most part adaptation to air-breathing to an early stop. many isopods and amphipods (Orchestia). km. The most important of these advantages is the abundance of available oxygen. is only 2. of oxygen are usually dissolved in a liter of water. attained by the pelagic different aquatic animals have at different times become few of these adjustments partially adjusted to an air-breathing life. with an average depth of_3795jn^ so far as investigated.5-70 m. as in tropical rain-forest. and as animals arc unable to raise themselves in the air. they are confined to a single layer. ice and desert are nearly or quite closed permanently to life. like those of snails and frogs. The surface of the oceans and inland waters combined amounts to about 361. whereas a liter of air contains 207 cc.000 sq.059. Such muscular activity as that of the wing muscles of insects. km. total inhabitable space available for terrestrial life is much smaller than that at the disposal of aquatic forms. without occasioning the complete transforma- tion of the mode of life. this makes possible an enormously increased rate of cojiibjistion for air-breathers if a sufficient food supply is provided.000 sq. Owing to the role played by oxygen in the release of chemical energy. Terrestrial life offers advantages that result in the luxuriance of such animals as are able to adapt themselves to it. in the common housefly. the terrestrial hermit crabs. are inhabited by living organisms in one stratum above the other from bottom to whereas the surface. In spite of these spatial restrictions.

which attained a high development before the appearance of terrestrial animals. many beetles and Hymenoptera. The advantages afforded by terrestrial life are counterbalanced by great disadvantages and dangers. is still large. earwigs. and include a few reptiles. the climbing perch Anabas. and among the arthropods. and favored the development of carnivores among made The earliest tetrapod vertebrates to enter the upon Amphibia. and all the lungfishes. cockroaches and grasshoppers. The number of herbivores among insects and myriapods. and the myriapods. insects. such as arachnids and vertebrates. and have undergone a renewed evolutionary development in consequence. the eel-like Amphipnons. tgrmites. Land plants. In spite of the low density of the air. which have called forth special . The gra/iTS. Another advantage gained by adoption of the terrestrial mode of life was the enormous amount of previously unavailable plant food. also. In view also favors arachnids unbroken extent of the atmosphere. are the pulmonate snails among the mollusks. mammals (bats). That shown lungfishes. many Diptcra. a few birds. a number of diverse types of animals have independently acquired the power of flight. were at first only a few competitors.52 Classification According to Environment lustrate this tendency.habitat The success of the herbivores in the terrestrial possible the entrance of carnivores. The apparatus for air-breathing among these fishes is diverse. feed especially upon mats of algae and of the attached diatoms. Saccobranchns (Siluridae). flight is the most perfect form of locomotion. and most Lepidoptera. which may perhaps be regarded as the Herbivorous insects include earliest terrestrial animals. Herbivores appear among vertebrates at a later stage in their phylogeny. afforded a food supply for which then. consisting of a pair of diverticula of the anterior of the part alimentary canal. air offers much less resistance to motion than water. are still largely insectivorous. as for example Misgurnus (Cobitidae). The invertebrate groups that have solved the problem of air-breathing. and a large proportion of insects and myriapods. and birds. some such mechanism made possible the development of the air-breathing vertebrates by the from the parent stock of crossopterygian fishes. Pulmonate snails. It has been mastered by insects. reptiles (in extinct forms). the mudskipper Periophthalmns. and the presence of the solid earth as a basis more rapid motion than is possible for aquatic animals. has proved successful. In consequence of its low density. are almost exclusively herbivorous. terrestrial mammals. an ecological division of snails. most true bugs. life.

which will must a have delicate but surface. will be subject to continued evaporation of body fluids and finally to the drying up of the skin and the entire permanently injured by drying. already developed in aquatic arthropods. would have been useless as a step toward terrestrial life without some protection to the breathing organs. but in this breathing existence. A respiratory apparatus. large permit the organ Two of carbon dioxide. much-branched imaginations of the body wall. skin is The glandular activity. a solid armor. be situated in the interior of the body. and it is covered outwardly by the epidermis. especially at night. The echinoderms have an external armor that serves as a muscle-supporting group the armor is formed by the deposition of lime in the subepidermal layers of the skin. ing apparatus is become impossible. no echinoderm has become adapted to a terrestrial or airskeleton. Thus. humidity Jiumidity reaches the maximum at only a few places and then usually only at special seasons. and groups of anirapid exchange oxygen mals. unshine. and secondarily as a protection. its cells are killed. and is usually far below the saturation point. conditions the rate of transpiration. The epithelium of the breath- The breathing especially subject to this danger. exoski'lcton overlies the outer surface of the body. of the air. the arthropods and the vertebrates. Most whose breathing organs take the form of thin-skinned. in the tropics. Crustaceans with external gills enclosed in a gill chamber by the lateral parts of the cephalothorax can survive in the air for short periods. their der adverse conditions. barometric pressure. and amount of sunSoft-skinned animals. in spite of the existence of a skeleton.Terrestrial Animals 53 The most important difficulty consists in the varying adaptations. The aquatic arthropods have ened outer layers of the skin. Those that have gone over . rate of wind. such as skin-breathing and sensory and body. The considerable protection against the drying of the skin. should crustaceans. formed by the thickThis armor serves primarily as a frame- work action for the insertion of muscles. is whereby the effectiveness of their This notably increased. which is thus unprotected against drying should the animal leave the water. to meet this situation most successfully. but only under favorable conditions of humidity such as are found at the seashore. Many hermit crabs and other decapods exemplify this de- gree of adaptation to life out of water. The humidity of the air. and important functions. combined with temperature. which is a vital condition for an effective protection against evaporation. find difficulty in acquiring such protection. were especially suited for terrestrial life by the structure of their skin.

Ic FIG. heart. /H. an genera of isopods. Among some Porcellio and Armadillidium. arachnids have evolved the so-called fan tracheae or tracheal bings. After Lang. The terrestrial jsppocls (Oniscoidea) are widespread. supplements the gills. or they have supplementary breathing organs. 7i. the epidermis of vertebrates is . tufts. respiratory cavity. crabs of some genera and land crabs (Gecarcinus. They usually occur in damp places. Such isopods predominate in dry situations. protection of the breathing organs is most complete in myriapods and insects. branchial blood canals leading rt. of their skin. The second or even all five pleopods may sometimes also be so modified. t'g. respiratory their en- pui. a\-a\. comparable* with the tracheal lung of spiders.54 Classification According to Environment permanently to terrestrial life and can thereby move far away from the water. with their tracheal systems. 4). with reduced gills (Fig. 4.. as in the cocoanut crab Birgus latro. as in internal breathing organ. which has enlarged inner surfaces of the gill chambers. pericardium. the trance into the pericardium. Vertebrates are protected against drying by the stratified structure and this is already developed in fishes.). acquire an invagination in the outer skin of their terminal branches. Independently of these. forming a much-subdivided breathing chamber visible externally as a "white body" (Fig. Cross-section through Birgus latro: Ic. re. g. which form a cover for the delicate gills. with a considerable number of genera and species. gills. branchial or lung cover. have apparatus for moistening the gills and for keeping them from sticking together in the air. where the gill apparatus on the underside of the* ab- P2 . such as penult. The first pair of abdominal legs. vessels leading to the heart. to the heart. The A less perfect tracheal system has also made possible an air-breathing existence in moist situations for the more annelid-like Peripatns and its allies. 5). lung or shell vessels leading from the heart. pulmonary same near domen is not in danger. p. In contrast with that of many invertebrate animals. etc.-J.

Outer brunch of tin- first abdominal appendage of the land isopod divided respiratory cavity. Of the various types of air-breathing or- gans developed among fishes. The cells die off with undergoes adaptive changes of horn and form a protective covering an accompanying development lor the remainder of the skin. These are composed of two conthird ) A vergeutly developed series that are evidently separate in origin. still single in formed of more numerous layers of small cells. is is increased. phylum. The gills of fishes. Pro- Fu. entirely . the land snails. by means of changes in the breathing apparatus. afford a sufficient surface only in the water. with much tection against evaporation in both terrestrial arthropods brates is based on structures already available. the Mollusca. has developed another large group of terrestrial forms. hermaphroditic forms without an operculum and sometimes. After Herold. These originate as a sac-like evagination of the anterior part of the alimentary canal. lungs proved most successful. and verte- Complete transition to the vertebrates. Arniadillidiuni nusutum.. 5. of layers of cells amphibians. JR . not unlike those of the higher Crustacea. as to an air-breathing existence became possible to the invertebrates.Terrestrial Animals 55 The outermost of these layers even among fishes. just posterior to the hindmost pair of gill clefts. and the horny stratum. The majority of the land snails in our zone are pulmonates. secondarily.terrestrial vertebrates the number composed of successive cellular layers. as their branches cohere in the air.

which is closed outwardly except for a relatively small opening and whose inner surface is only epithelial layer The slightly increased by the projecting walls of the blood vessels. or a leal. for example) against evaporation. in which the entire body is covered by the protective layers. Other animals that have accomplished the transition to land life . content of it possible for pulmonate snails to the air makes oxygen exist with a reduced breathing surface. 10 which reduces the loss of water by evaporation. in spite of the fact that the shell forms so favorable a protection against evaporation. Many pulmonate snails have lost their shells in the course of their evolution. The gland cells are superficial in the prosobranchiates. Like the crustaceans. others adhere of the foot to a stone. the slugs have gained a more slender form and are thus better able which they find protection Limax maximus. but in correlation with its small extent their activity and energy development are scarcely greater than those of marine snails. Although it is usually possible for the snail to withdraw entirely into its shell. Snails may pass through periods of drought by closing the shell with the operculum. snails with separate sexes. Their mucus (in is often tougher than in the shell-bearing snails and affords a more to avail themselves of existing retreats in effective protection. and this naked part of the animal is protected against evaporation primarily by a coat of mucus. the shell. Toward the tropics land snails of the prosobranchiate become These are operculate group progressively more numerous. ^ Land snails differ their aquatic relatives in the position and number of their skin glands. when an operculum is present. In dry regions snails usually come out of their shell only during the fall of dew at night or during and after rains. However. a part of its body must be stretched forth for creeping and other activity. a tree trunk. in the pulmonates they are sunk deeper into the sub- by means from and open externally only by means of narrow canals. by this loss and by the smooth- ing out of the visceral sac. within the epidermis.56 without a Classification According to Environment shell. which was formerly contained in the shell. breathing surface of the lung is notably smaller than that of the muchThe richer feathered gills of the marine water-breathing snails. Land snails arc thus radically distinguished from arthropods and vertebrates. the non-oper- culates may secrete a film of mucus that serves the same purpose. Their breathing apparatus is formed by the mantle chamber. the aquatic gastropods possess a protective structure. as a preadaptation to protection against evaporation for the forms taking up terrestrial life.

Protection against desiccation is extremely unequal in the various groups. The thicker their shell and the greater their capacity for existing in a state of suspended animation without food. All these are found. earthworms. are less subject to evaporation on account of having fewer glands. too. such as land planarians. in so far as they are not protected by a dermal armor. earthworms. and the air-breathing crustaceans. mesica and jxeric forms on the basis of the extent of their tolerance of aridity. against injury by drought by means of special adjustments that enable them to spread into dry regions with only occasional humid periods. such as Helix (Eremina) descrtorum. include -land planarians. These animals can time without sign of life. and. It has been noted above that excrete the water taken up by their skin through their kidneys frogs and collect it in their bladders. Even though a sharp line cannot be drawn between them. 7a exist for great periods of amphibia belong to the hygric are insured these. and a number of leeches. by the production of mucus in their skins. Hygric_aniiiials. 20 The air-breathing snails also be- long to this category. Thus in Ger- many. in situations with very humid air. Some are able to save up water Of the terrestrial vertebrates. One may find every transition.Terrestrial are a Animals 57 few forms with incomplete protection from evaporation. from aquatic animals that can exist for a short time out of their native element to forms that live continuously under conditions of extreme dryness without drying out. and their capacity for life in the air is equally variable. This water contains only very small in their . such as Armadillidium and Porcellio pictus. but they can penetrate into relatively dry regions if the air reaches the necessary degree of humidity from time to time. have been known to revive after more than four years of suspended animation. with few exceptions. Pcripatus. group. They protect themselves against temporary drought by retreating into holes in the earth or beneath stones or logs or similar objects. steppe and desert snails. the more possible it becomes for them to inhabit arid situations. in a state of aestivation. While the German grapevine snail Helix ponwtia can live at most a year in such a condition. Helix (Xcrophila) ericetorum lives on dry and sun-burned slopes. land leeches. that live in drier places. and elsewhere desert snails extend as far into the steppe and desert as does the occasional deposit of dew. as they are able to retire into their shells and close them off during the dry periods. ncmatodcs. Isopods. the But bladder for times of drought. it seems useful to classify air-breathers into hyjjric. They are able to come out of their shells only in moist air.

of boring bettles (Anobium). The capacity gradations. at night. !fi Most insects are creatures of the sunlight. so that smaller amounts of blood can care for the distribution of nourish- ment and water for the removal of excreta. am! arachnids and on the other by reptiles. as a part of the excretory products is stored in the fat-body. Finally. as before rainstorms. Many relatively soft-skinned insects avoid direct sunlight and are active only in moist air. These insects are able to use metabolic water and can further conserve their water supply ' by excreting form. their nitrogenous wastes in solid rather than in liquid Many in habits. however. young mole crickets. in the twilight. insects. the tracheal breathing relieves the blood of an important function.58 Classification is According to Environment almost pure. These groups are by no means excluded from existence in humid regions. are unfavorably affected by higher humidities and amounts of rainfall. as in steppe animals like the camel. their whole body is swollen by the distended bladder. How in the small the need for free may be in insects is best shown forms that can live on very dry food. Some. mosquitoes. Xeric animals are represented on the one hand by myriapods. birds. to withstand dry air without danger appears in all Many myriapods such as Lithobitis are inadequately pro- or after dewfall or rain. They are protected against loss of water by their dense body covering and have few or no skin glands. when the frogs bury themselves after the close of the short rainy season. and amounts of urea and they survive twelve and even eighteen dry months in this condition. which live in wool and hair. Since humid regions and situations are relatively few. the xerocoles in general have a wider distribution than the hygrocoles. 16). by carrying the oxygen directly to the points of consumption. such as the larvae of meal worms (Tencbrio. or of the skin beetles (Atlagcntis) and clothes moths (Tineola). Nor does the excretion of waste require much water. These include mayflies and stoneflics. which eat the dry wood of old beams and old furniture. tected against evaporation. . which may easily be killed by the sun's rays. which swarm only in moist air and otherwise remain in hiding in places sheltered from the sun and wind. probably in consequence of specific adaptation to arid conditions. Thus in the Australian deserts. and most termites. and mammals. sec p. desert-inhabiting animals may be cryptic and nocturnal and consequently actually hygric rather than xeric. which carry on their building activities and foraging expeditions only at night or in humid weather. There is a dearth of exact information about the desert toads and spadefoot toads of North America with respect to the physiological aspects of their aestivation.

When they are present in abundance. but as crystalline uric acid. in the desert he may sweat the rate of a liter per hour. aardvarks. are also diverse in this respect. but our knowledge of them is as yet inadequate. part in the regulation of body temperature by giving off large amounts In dry seasons. excreting urea in solution. for the most part with a small number of skin glands. without noticeable perspiration. and horses. apes. which promote evaporation. The impor- may endanger mammals can hold out the life for tance of these relations in the geographical distribution of vertebrates has not been sufficiently studied.e. Between these extremes are such forms as the hedgehog and squirrel. by skin glands and contrast. give their excrement. getting along with the water taken with their food and with the water of metabolism. They exist in even moderately dry regions only by means of special physiological and ecological adjustments. concentration of the urine. and skin glands are very unequally developed in the various mammals. skin protected only by a thin horny as specifically hygric.5 liters of water per day through his skin When by insensible evaporation.. as in men. the loss of water is naturally larger than in forms in which they are nearly or entirely absent. and mammals. and many antelopes and gazelles. they economize on water because their excreta are not their evacuated in a dissolved state. ' 1 1 1 man requires 5 to 8 liters of water daily. off a good deal of water. many months without drinking. i. a of water. the losses of water are notably larger. the soluThe ostrich is an exception waters are reabsorbed in the kidneys. reptiles. birds. 10 At at high temperatures and in dry air the lack of water for more than 24 hours of a man. It must be admitted that regulative . or even three times as much. much They may be contrasted better designed to withstand dryness than as water-savers and water-spenders." as do mice and porcupines. " and thus take skin glands are transformed into genuine sweat glands. and the carnivores. with skin glands sparingly scattered over the body. tion. through their There are great individual differ- ences. The three remaining classes.Terrestrial Animals 59 The Amphibia with their glandular layer were characterized above air-breathing vertebrates are xeric in very different degrees. as in most rodents and certain ruminants. Besides this. and water content of the excrement are important factors in water economy. hyraces.' 5 Mammals. Reptiles and birds arc in general are most mammals. A man gives off more than 1. On the other hand. in Reptiles and birds have an advantage over mammals complete lack of skin glands. Evaporation through the lungs.

Kaempfferia kacrnpffcri. arthropods. which they The soft-bodied terrestrial forms are compelled to rest the whole extent of their bodies on the earth. is a dwarf.000 kg. when greater degree of independent motion is possible only an internal or external skeleton is well developed. the air does not as- supporting the body as the water does. for live furnishes support. the loading of the less resistance than water. Domestic cattle. is only one third as great. though the bison of North America formed an exception. and vertebrates fall behind their water-breathing relatives in maximum size. as the land planarians. long) and some Palophus. doubling the size of the body will require more than a twofold increase in the strength of the supports. in. in weight. Hence the air-breathing. This necessitates a general stiffening of supporting structures. their salivadripping mouths. or the giant crab Compared with the blue whale of up to 30 Homams. This Thus cattle. limbs limits the size of the body. but these too are firmer in structure than their aquatic relatives. have become adapted The physical character of the air as a surrounding medium condi- tions certain peculiarities of structure in terrestrial animals. such as water buffaloes. long) are much smaller than the rock lobster Palinurns. group of animals is little represented in steppe regions.. and even the weight of the giant extinct saurian Brontosaurus. except the Mycetozoa. the body offers less friction when in motion. are great expenders of water. earthworms. m. Even the largest land snail of the genus Achatina does not reach the measurements of the marine Tritoniwn. skeleton-bearing forms of mollusks. (Dynastes hcrculcs. and snails demonstrate. Forms \Vith a gelatinous in body are excluded from which the decaying wood terrestrial in life.5 mated at 38. are swamp dwellers.g. The giant insects and arachnids such as the Hercules beetle grasshoppers the lobster (e. long. since in consequence of the laws of statics. ever.000 kg. may take place within the same with their numerous skin glands. howto going without water for several days 10 at a time in the arid peninsula of Lower California. 2. and their soft excrement. 15 cm. which is estiin length 3. and certain features that were excluded by the nature of the aquatic habitat become sist 11 possible. long and 4000 kg.60 Classification According to Environment adaptation to special conditions species. A much Raised from the earth by means of stiffened limbs. and many forms. and a weight of as much as 108.. and as a consequence most of them live in humid regions. The terrestrial ani- . and at the same time the air offers On the other hand.. On account of its low density.5-30 cm. the elephant. 20 m.

the one is stimulated by liquid substances. density of the air is also accountable for the fact that (parasites aside) there are scarcely any sessile terrestrial animals. the other by gaseous ones. By this division of labor. which aquatic animals and normally resort Internal fertilization may still behave water at breeding time. is conceivable. are adjusted for distance near objects. but it is in- to variable in the fully terrestrial ones. the random transport of male sex cells to the eggs by means of air currents. take place in aquatic animals. The fertilization of the complished by means materials on the part sion of of a copulation. of the males. are less different in size from their aquatic relatives. the sessile habit supply. higher development of the organs of sense is a general accompaniment of the transition to terrestrial life. in contrast with aquatic animals. To be in successful. and are actively focused for and fishes. . which is so common a process in aquatic animals. i. In squids The chemical senses of aquatic animals are developed in the airbreathers into the senses of taste and smell. among which sessile forms are com- The low mon. as eggs is usually acThis means a great saving in free emis- compared with the sperm into the water. but this possibility has never been realized. as in some Peripatus and in the viviparous mammals. such as the land planarians and earthworms. importance.. which present faint and indefinite images in water. Fertilization is internal in all air-breathing forms with the exception in this respect of frogs like and certain salamanders. when at rest. the eye increases in A evolved. The lack of a plentiful supply of air-borne food particles also contributes to the necessity animals.e. An arrangement like that of wind-fertilized plants. but the air can bear larger particles only in the less frequent instances when it is in rapid motion. since the delicate larvae hatching from small-yolked eggs would not be able to feed themselves. since small animals suspended depends on an abundant food and organic particles may be passively the water and be brought to the animal by the creation of a current. On account of the greater transparency of the air as compared with that of the water.Terrestrial Animals 61 mals without a skeleton. though no new types of eyes have been In terrestrial vertebrates eyes capable of accommodation when at rest. the eyes remain adjusted for near objects and accommodate actively for the more distant ones. Only when care of the young takes place and the emfor large-yolked eggs among terrestrial bryos are supplied with food in the mother's body can the eggs be small and poor in yolk.

within certain limits. this makes it possible for them to live in zones and at seasons when other animals are dormant on account of cold. Among primarily . acceleration of nerve impulses. whales. This importance of smell depends in part on the tendency toward uniform diffusion of gases and the resulting wide and uniform distribution of odorous substances in the air. however. work together to raise terrestrial animals to a level and more diversified behavior than that of This aquatic animals. in inanimals. It is of special consequence to nocturnal and cave animals. As the organs of taste and smell must have cells of living protoplasm at the In the surface. as compared with that of water. Organs of hearing are enormously more developed among terThese are found in terrestrial vertebrates. and the cries of parrots lead monkeys from The much animals afar to isolated trees with ripe fruit. Warm-blooded animals are rendered independent of external temperatures. Some steppe mammals catch the scent of freshly fallen rain many kilometers away. are vertebrates protected by being placed in speair-breathing they cial chambers and are there kept moist by means of In aquatic animals. vultures see their carrion from an extraordinarily great distance. and seals. has made among terrestrial forms. They are wanting in primarily restrial aquatic animals except in fishes. Sense impressions at great distances. they are especially subject to the danger of drying. shortening of the latent period in muscle contraction. ^ Uniform optimal internal temperature produces. in addition. possible the appearance of homoiothermal It is true that some homoiothermal animals are found in the seapenguins. perhaps among arachnids. the organs of chemical sense may be distributed over the whole surface.62 Classification According to Environment which. and sects. are not possible for aquatic animals. even in fishes. males of many Lepidoptera are attracted by the odor of the females from distances of several kilometers. all of which result in an > All these factors of intensification of life processes. lower conductivity of the air for heat. higher level also finds expression in the frequent development of a more complex central nervous system and in more intense activity the attendant phenomena of instinct and reason. must become dissolved to be perceived. The as in terrestrial forms. for examplebut they are secondarily aquatic forms with especially well- developed insulation. and acceleration of digestion. cavities or glands. The sense of smell acquires a great importance in some air-breathers (many insects and most mammals) for general orientation and may even replace vision in importance.

birds. such as appears among insects. The differences in humidity are. pressive. there is a much greater diversity and variability in the conditions to which terrestrial animals are subject. 23 Of arthropods. although the sea is its original home. and the genera and for the 15 This is even part even the families of the Jurassic are extinct. ^This explains the much greater number of species of terrestrial as compared with and aquatic animals remarked upon above. finally. excluded in the aquatic habitat. cirripedes. ~The genera of fishes have changed relatively little since the Cretaceous. soil ences in the nature of the of topographic relief. which originated in the Silurian period. especially in the sea. many genera of Crustacea can be traced back to the Mesozoic. and fishes are in any degree comparable to land animals along this line. is more imlife. In the sea temperature variation has an amplitude of only 26. Nephrops. teleosts like . oldest genera of animals now living are aquatic forms witness the brachiopod Lingula (Cambrian to Recent) and the gastropod animals and the shorter duration of the The Pleurototmiria. of course. and social life. and only 20 at any one place. 43 In contrast with these. and mammals. All sharp these factors together result in more diversity and complexity of adaptations and of directions of evolution. is developed among the primarily aquatic forms. more notable among the vertebrates. ScalpelAmong decapods Callianassa also appears known from and Palinurus. the genera of most insects range only into the Tertiary. It also explains the fact that genera of aquatic animals extend so much farther back in geologic time.Terrestrial Animals 63 aquatic animals. only cephalopods. Ceratodus known from the Triassic is very close to the living Neoceratodus. In contrast with the great uniformity of habitat conditions in the water. many modern genera of selachians are known from the Jurassic and Cretaceous. and favor to a high degree the transformation of forms both in space and time. crustaceans. deeply of affect its animal on account climatic The great differThe influence results. Pollicipes appears in the Jurassic. and probably Homarus are the Cretaceous. Thus. in the strict sense. The origins of our present-day land snails date from the Eocene. On land the temperature may go far below zero and may rise to more than 70 C. those of the fresh- water mollusks from far back in the Cretaceous. the separation of land areas by water stands in contrast with the much greater continuity of the oceans. Among modern him in the Jurassic. which is a correlative of the more rapid evolution among land ^ life of a species among them. in the Cretaceous. animal life reaches its highest less perfectly development on land.

either for their entire life or at least for their larval period. together with some spiders. contrariwise. and sirenians. In fresh water. air. and the genera of mammals rarely extend as far back as the Miocene. great numbers of insects have taken up their existence. such as auks and penguins. as in the sea snakes and aquatic ondarily sea turtles. which thus become secondarily aquatic animals. sea otters. of Lake Geneva and from the waters Limnaca witness deeper stage. Marine arachnids are limited in number. there is a superficial network of blood capillaries in the phosis. such as the sea turtles and sea snakes. Sometimes. Only the water mites and insect larvae with tracheal gills reacquire the Others are in the transition ability to obtain oxygen from the water. and However. whales. the water affords a new sphere of action for air-breathers in which they have no competitors on their own evolutionary level. and many pulmonate snails. all hydrachnicls. seals. numerous birds. whales. Thanks to the advantageous characters acquired in connection with terrestrial life. and many other In contrast with these. often far from land. the reptiles have undergone great changes since the Cretaceous. The fresh-water inhabitants also include a number of reptiles (crocodiles. One small family of water striders. the Halobatidae. various mammals. Upper Cretaceous. arid in the axolotl and the perennibranchiate salamanders they reach sexual maturity without metamor- perhaps a question as to whether these larvae are secin a strict sense. and even such thoroughly aquatic mammals as seals seek the land at the breeding period. This explains the readaptation to life in the water by terrestrial forms. In the ocean it is chiefly the vertebrates that have returned to aquatic life.64 Classification According to Environment in the Clupea and Beryx appear existing genera in the Eocene. turtles. It is mouth and on the jaws. spend" their whole life on the surface of the sea. is which is the chief source of advantage for the terrestrial always retained in the secondarily aquatic vertebrates. the breath- ing of forms. and almost no insects are truly marine. Crocodiles and turtles lay their eggs on the shore. such as the otter and beaver. the aquatic turtles with accessory anal breathing organs. With the exception of a few viviparous animals such v as sea snakes. The larvae of amphibians are gill breathers. and in part homoiothermal animals. and sirenians. The dipterous family Chironomidae has a few larvae that live below the tidal zone. v . the secondarily aquatic vertebrates abcwe the Amphibia retain their terrestrial breeding habits. the penguins lay and brood their eggs on land. in part poikilothermal forms. and numerous snakes). in the warmer seas.

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. 1907. Sollas. Macinillan: viii. 1 . Textbook of paleontology. Ges. 43. W. 7:1-16. von. Ergeb. Max. Zuel/er. K. 1. 30:163. 28 figs. "On the origin of fresh-water faunae. 1884. A. Weber. don. Relse Nederl. 1932. 600 and 664 ff. 1476 figs. 42." Nature. 67 Homer W. 1907:90-94. Margarete." Quart. "Water regulation and its evolution in the fishes.. 41. a study in evolution. "Die Siisswasser Crustacecn des Indischen Archipels nebst Bemerkungcn iiher die Siisswasserfauna im Allgemeinen. vol. pi.)." Zool. Ost-lndien. naturforsch.. (pp. 706. 1892. English ed. Freunde Berlin. Rev. 44. LonZitti'l. 1900.. J.Bibliography 40. 2:528-571. 30. "Ueber den Einfluss des Meerwassers auf die pulsirrende Vaeiiolc /* Sitzber. Biol. 574. Smith..

After only three years the thickly covered by blue-green algae. 11 species of ferns and 15 flowering plants were found. bugs. 1883.Ba triers to Distribution and Means of Dispersal THE EXTREMELY RAPID REPRODUCTION OF LIVING ORGANISMS CAUSES THEM to spread in every direction. A visit in of 1908. and moths. Animals had the in followed Even 1889 there was a whole list plants. and 3 mammals (2 bats and Rattus rattus). east of Java. thereby being prepared for the advent of higher plants. the small volcanic island of Krakatoa. was with 12 ferns and 50 flowering plants established. which 240 were arthropods. and the composition of this flora was notably different from the earlier ones. 41 km. In 1906 a new examination yielded 114 species of plants. a species of earthworm. and animals present could have arrived by air transport. among which were one snake (Python rctictilatus) 26 breeding birds. beetles. promptly of arthropods: spiders. flies. 25 years after the eruption. Investigations in 1920-1921 yielded 573 species of animals. 2 species of reptiles and 16 birds composed the vertebrate element. A visit in 1897 showed further progress. Comparison with neighboring . butterflies. soil perhaps. which destroyed it in part and covered the remainder so thickly with ash and pumice that no plant or animal was left except. constantly tending to" enlarge the area which they inhabit. so that no place in any way capable of supporting life When life remains unoccupied. a catastrophe such as a flood or a volcanic eruption destroys at some place. shows that the fauna then had about 60 per cent of the ex12 Approximately 90 per cent of the plants pected number of species. 4 species of land snails were found. yielded a collection of 263 species. it is soon replaced. The rats and islands 68 . was the scene of a tremendous volcanic explosion. and even a species of lizard (Varanus salvator) was present. On August 26.

however. families. They are conditioned by and change according to the ecological valence and the means of dispersal. The rat present was represented by 2 subspecies and therefore probably came from two different sources.^ecies. Sibesia. possible distribution of plants and animals. according to the classes and orders. There were 36 species of birds. and those that do exist are is more or less intangible." The tions. by physical organic environment and by the constitution of the organisms themselves. These highways are not common to all animals. ^ ' - r. Barriers anii mtenis of dispersal for marine animal life single vast continuous mass of water. Barriers to dispersal are fewest for marine animals. The possible distribution of existing species accordingly depends upon the bar- and organisms the capacity for dispersal Definite limits are set upon the means of dispersal at the command of the orand on its vitality and adaptability. could have swum. is not unDispersal does not take place with equal success in all direcin different is also very the and unequal. Barriers and means of dispersal are fundamentally different for aquatic and terrestrial animals. rotifers.5 km. limited. aquatic animals. fl( &? kf~ ***"M'< *f * '< "*' * '-. The oceans form a A Mediterranean and Baltic seas. and tardigrades were found on Krakatoa in 1928. and among the riers present. A world-wide distribution among mammals found only among some marine whales. iu question. the 47 species of vertebrates on the island included 3 species of bats. and perhaps the python. so that these divitially separated have no few basins. and well-marked highways may be found by which dispersal has taken place. in addition to the python previously recorded. 5 of lizards. so that the new inhabitants must have come from at least that distance by means of wind and waves and other agencies of "fortuitous dispersal. and a crocodile. with interr^J^iiAtJI*pttlepgeog^aB * n y an ^ evolution. and lug^--'^aMvM^ generally involves the historical factors. 03 Rhizopods.flE^felations tend to be different for marine forms and for ftasKf^Mlfc. The crocodile. is 18. and even to the genera and species to which the animals belong. The nearest island not destroyed by the eruption. away. such as the sperm and the whales. are connected with the open ocean only by narrow straits and are thus rather well delimited. killer .Barriers and Means of Dispersal 69 lizards probably came on driftwood or boats. The distribution of genera. such as the sions sharply defined limits. 45 years after the disaster. 23 By that date. Thus definite ganism of distribution patterns appear. iu tuiB. only parinto divisions by the continents.

rarely is above 3 C. The greater oceanic depths form a barrier to the spread of such animals of the littoral region as are confined within a narrow variation of depth. They can migrate only along the coast. and some of them are accordingly found in all seas. in themselves. The Indian and Pacific oceans are continuous in the warm zone. irregularities in the topography of the ocean floor. and channels. In consequence. and a great many wide range and differ strikingly from the fauna Of the 160 species of the genus of crabs Cycloin the Red Sea. pass around the The continents to the north or south unless prevented by some other factor. 18 and a few of the Metazoa is common to the east deep sea. Such animals may be called stenobathic. 31 Cold. The salinity of ocean water. Strong swimmers and other widely distributed pelagic forms. they afford no hindrance to the spread of pelagic forms. and the abovementioned whales. only two occur to the west. and below 2000 m. even when the land masses arc narrow. varies only between narrow limits. and are consequently less saline. can. species of the marine of tropical Africa. If they meet a thermal barrier. the whale shark. the Atlantic is separated from them by colder waters.70 Barriers and Means of Dispersal Land masses inserted between the oceans naturally form insurmountable barriers to aquatic animals. from the east African coast through the entire Indian Ocean and far into tropical Polynesia. With the exception of a few pelagic animals.. 40 united by the cold waters of the oceanic depths. eurythermal pelagic forms by no barriers. species have this of the Atlantic. of course. like the Isthmus of Panama or of Suez. Temperature sets a limit to the distribution of many stenothermal marine animals. are theless much less may form effective barriers in marked than on some places land. the uniformity of the marine life. but neverfor the bottom- inhabiting animals although. deeps.and depthtolerant animals have a wholly continuous habitat. so that the cold oceans of the two poles are brates. scarcely a and west coasts On the other hand. while the others metopa extend for the most part far into the Pacific. even in the tropics. their distribution may be limited. is very striking among both fishes and inverte- Most genera of mollusks. Such forms include the schizopod crustaceans Euphausia pellucida and Eucopia australis. many inhabitants of the are restrained . Where arms of the sea have an abundant inflow of fresh water. for example. in general. The warm parts of the oceans are separated from each other by cold regions at the poles. The temperature drops rapidly with increasing depth. such as ridges. a few of brackish water.

below the surface. either completely sessile on the bottom like the corals. water near the mouths of large rivers may be a barrier to the spread of littoral forms in shallow waters. many animals are restricted in position. with a depth of about counterpart. but it may extend deeper than 150 in. or oysters. The greatest degree of freedom is enjoyed by those that can rise freely in the water. its relatively shallow. or very limited in their power of movement. like sea anemones and some mollusks. are these creep. currents are of especial importance to the distribution of sessile marine forms that have a free-swimming larval period. But even in the water. Much plankton is thus transported by the Gulf Stream from the warmer parts of the Atlantic to the coasts of Europe and even to the neighborhood of Spit/bergen. All are deon movements within the water or on the movement of the pendent water itself.' 100 in. The oceanic Littoral animals is may be carried in this way to places where the depth too great for them to develop further. 19 movement of the water operates to distribute the weak swimmers and especially the plankton. . is also somewhat shallow. vertical movement. Passive dispersal of marine animals is almost confined to the surface forms. The eggs and larvae of many animals of the depths rise to the surface. which walk or run with restricted to the bottom. the suspension or by active swimming. The suspended plankton is dependent on passive dispersal by currents. the fewer are the effective barriers. bryo/oans. for terrestrial forms. whether in greater the capacity for swimming. the Kuroshio of the West Pacific. For example. reaching down about 700 in. may become subject to its motion. none of the species of of sea urchins of the Patagonian coast are La Plata. which north of Cape Hatteras is known 800 is as the Gulf Stream. 45 found north of the mouth There are fewer methods of distribution for marine animals than There are no true flying forms. lies in the westward-bound north equatorial current in the Most wave motion tends to be negligible below about Atlantic. Such animals may escape the current by downward of 3000 m. which is composed of plants and animals that do not swim independently of the motion of the water. and so be destroyed. By contrast.Barriers and Means of Dispersal 71 The freshening of the surface neighboring ocean are excluded. and storm waves have This type of disturbed sand nearly 400 in. The Florida current. Many marine animals such as worms or snails. 5 '' or. Similarly. The majority of have free-swimming larval stages. which more or less speed. and crabs. by rising. in. The level at no motion varies and.. although usually nearer the surface..

which varies greatly among different forms. and even reef corals (Pocillopora) have been found attached to a floating piece of pumice. are widely distributed because they all Almost with exceptional powers of suspension. The larval life of the trochophore of annelids and mollusks seems to be much shorter (4-5 days). only Discina atlantica This is explained on one hand by the fact that this species free-swimming larvae to the surface and on the other by the is distinctively a form of the greater depths. the phosing. GT Transport by other marine animals may also occur. so that the mature larvae find suitable habitat conditions almost wherever 4 they sink to the bottom. whales and to sharks and other fishes by disks. The ten species of sucking fishes (Echcneis. Among has a world-wide distribution. and Sqnilla cruposa on the 7 coasts of North America and Africa. Many crustaceans (Euphausidae. the Crustacea of the group Stomatopoda. Conodactylus chiragra occurs in all oceans. Remora). The distances traversed depend upon the rate of movement of the current in question and the duration of the free larval period. The Tcrebratulina zoea stage of the decapods continues for 25 to 30 days. Many marine forms Thus the American sea anemone New Haven and Boston and thence reached Europe. are distributed to all tropical which attach themselves to means of their large sucking seas and warm by this means. in 1896 and at Busum in 1920. rise of its the brachiopods. Barnacles ( Balanus and Lepas) are found on driftwood and on actively swimming Even the adult stages of sessile forms may be animals including whales. and even sea snakes. whose adults arc slow-moving bottom dwellers. turtles. 50 Plymouth . have a long larval life transported by currents if they attach themselves to a moving object. Sagarfia luciae was carried along the coast from the south to arriving at are slight. Penaeidae) and deep-sea fishes (Muraenidae and Scope36 This procedure is naturally effective for lidae) afford examples. and echinoderm larvae may drift for periods of 20 to 60 days before metamorseptentrional is itself after attaches larva of the brachiopod 10 to 12 days. the distribution of these forms because the larvae come within the influence of the currents. though are their own swimming powers transported by shipping. The bivalve mollusk Dreissena attaches itself to wood by its byssus.72 Barriers and Means of Dispersal undergoing their development in the light. as does that of the planula larva 10 of corals and sea anemones and the Miillerian larva of turbellarians.

which are absent from the Danube. but migration from one river system to another is not facilitated by this means. and Pcrca volgcnsis. Thus in New York. yet to go from one to the other by water would require traversing thousands of miles. Remarkably long distances by water connect portions of different drainage basins separated by only a few miles of land. Rivers for the most part flow into the sea. tends to prevent a mixing of the salmon populations of adjacent river systems. Thus the Danube basin is distinguished from that of the Rhine by the difference in the migrating fishes. the species are distinct. difThe homing instinct of ferentiate the Danubian fauna still more. with different species of sturgeon and salmon. which are absent in the Rhine. which lie eight miles from each other. in the little or no Danube.Barriers Barriers and Means of Dispersal 73 and means of dispersal for fresh-water animals Conditions dispersal are decidedly less favorable for fresh-water animals than for marine forms.* and the presence of eel and shad in the Rhine. River systems are sometimes connected because one stream pirates the headwaters of another. would have been even longer. Nearly all the genera of freshwater fishes of Africa are different from those of South America. Before the opening 1 of the Chicago Drainage Canal in 1900. are common to the two sides of the Atlantic. Neighboring river systems are often decidedly different in their fauna on this account. All permanent freshfor direct water basins are separated by land of greater or less extent. and the watersheds then form the dividing lines between such faunae. al- though the Atlantic breeding grounds of the European and American eels overlap. which brings them back to breed in the streams of their nativity after their sojourn in the sea. A. almost half of which would be through salt water. sturio and S. especially if they flow into different oceans. The absence 1 presence there of of the sticklebacks (Gastcrostcns) in the Danube and the numerous eastern fishes such as Abramis sapa. Conversely. Fresh-water animals that can enter the sea may thereby have a very wide range. salar in . since the sea offers as effective a barrier to most fresh-water forms as does the land. and " only those like Aritis whose representatives enter the sea. Dispersal within a body of quiet water meets with Acipenscr ruthcnus and Salnw hitcho the Rhine. Gobio uranoscopiis. Lake Chautauqua of the Mississippi River drainage is about 125 miles from Lake Seneca of the Atlantic drainage. salmon. the water journey from Lake Chautauqua to Lake Erie.

living on them for a short time as parasites. and tongues of swimming and wading birds. ginning 'J Passive dispersal plays an important role in inland waters. maintain itself on the bottom. bottom forms like snails and eels or powerful swimmers like the Salmonidae are able to cope with a current most easily. which are the only active swimmers in fresh water. Such has confirmed by direct observation for rhi/opods. winter eggs of cladocerans. where they establish themselves. frequently serve other animals as a means of transport. and then attach itself to another fish. is The salmon Venern or in the streams draining into it. feet. without losing its vitality. The marine fishes that enter the rivers of Malaysia bring their paraThus Rosinella tyjnts. 69 The passive distribution of fresh-water animals by Hying forms is more frequent and effective. threadworms. bills. Frog spawn may be kept in the air for considerable periods (up to four days). the current has an important influence. Birds are the principal transporting Eggs and forms in a dormant state become attached to the agents. Suspended animals are af- fected by even a slight current. if they are not too large. during which period they are transported by their hosts. In strong currents upstream dispersal is made difficult.74 hindrance. and eggs of snails. either in Lake barriers to dispersal. thus in the Havel lakes near Berlin the lower lakes have a larger number of the cladoceran Bosmina corcgoni than the upper. Thus the larvae of river and pond mussels (Unio and Anodonta) clamp themselves to the fins or gills of various species of fishes. since they can spread with the current but not against it/' 5 Waterfalls and rapids accordingly form well-marked unable to pass the falls of the Rhine and is therefore absent from Lake Constance. of the nineteenth century this condition was changed. and the ported Sea of Sinkarah. because the young eels with the building of locks at the bewere unable to pass the fall. for a time This isopod is known to leave the fish. 3 " Transport of tiny animals from one body of water may also take of ducks' feet . the Capuas River in Borneo. is resitic isopods with them. webs Bivalves occasionally clamp themselves to the and may be transported. The isopods of the family Bopyridae parasitize marine crustaceans (Palaemon and others) and are carried into rivers by their hosts. if the weather is cool and 35 The conditions for its transport are damp. a cymothoid. Fishes. Above the Trollhattanfall in Sweden there were formerly no eels. been transport statoblasts of bryozoans. from Gulf of the Bengal. rotifers. Barriers and Means of Dispersal In running water. feathers. therefore favorable. which may be a fresh-water species.

where ice may connect various islands. Gudger "rains of fishes. rivers often have more importance as a connection . Snails of the family Ancylidae have several times been seen adhering to the wing covers of beetles. has repeatedly been found with the small bivalve Sphaerium clamped to its legs. Eurasia and Africa. wide.* and Piaidium has been found attached to a water bug. The small ostracod 74 Such Cyclocypris laevis is transported in this way by flying insects. Great streams like the Amazon and its tributaries may limit the range of many forms of forest mammals. entire south bank from the Atlantic to the Andes and has not become established on the north side of the river. however. In addition. These separations are bridged at only a few places in the polar seas. 2 * animals. means of passive transportation are of little consequence to marine animals. effectively limit the dispersal of animals but more freis quently have only minor significance as barriers. there are a great number of islands of all sizes. In general. but they are important to fresh-water animals. 12 The La Plata estuary forms the southern limit of the range of the capybara and the northern limit for the viscacha. the Americas. a separation by the sea conditions the distribution of animals. Sakhalin and the polar islands of North America have thus been connected with the mainland. The larval stages of parasitize aquatic beetles and bugs and are carried away by them on their aerial journeys. frogs. the principal land masses are separated by sea water into three hugh blocks.Barriers and Means of Distribution 75 hydrachnids place through the agency of insects. on account of the discontinuity of theirs. salamanders. The ant bird Pldcgopsis It occurs along the nigromactdata furnishes a dramatic example. and means of distribution for terrestrial animals animals. however. Mozambique Channel between Madagascar and which Rivers may only 400 km." has given an excellent resume of authentic Other animals such as tadpoles. 1! * and mollusks are transported Barriers in this way. with their continuous habitat. as does the Africa. Fresh-water animals may be carried up by tornadoes and deposited at a distance. birds. The oceans are thus the principal barriers to the free dispersal of land animals. Harriers to distribution are of especial importance for terrestrial Whereas the oceans are connected. 28 Dytiscus margitudis. and Australia. citellus. and narrow straits may separate very distinct faunae. and butterflies. 37 The south Russian Dnieper separates ranges For lower of the spermophiles Citcllus suslica and C. a water beetle with strong flight. entirely separated from other land masses.

Sumatra is traversed for its entire length by a mountain range that separates a northeastern and a for certain species or southwestern coastal strip. have an unusually wide range both from east to west and from north to south. in a region in and subspecies of land animals. importance Mountains and deserts as barriers. river. if the is not too broad. and even a great number of species. in some respects they are even more effective. has completely changed the lower part of its course nine times in the past 2500 years. The faunae on the north and south slopes of the Himalayas or also prevents on the east and west sides of the American Cordillera are quite disThe fauna of Africa south of the Sahara exhibits a striking uniformity. The Hwang Ho. but it had not crossed the mountains and was absent on the east coast in 1917. insects. 8 * 14 ' 6 * Mountains of even moderate height may form effective barriers even for whole faunae. introduced into New Caledonia in the 1870's. land isopods. or of the Ganges. partly because of intimately associated climatic changes. has increased on the west coast to such a degree as to be a pest. Floods cause shifting of the stream-bed and create and Pieces of woodland may be passively transferred from one bank to the other. mammals. many animals from reaching or crossing its passes on account of the reduced temperature and lowered atmospheric pressure. The sea. and tinct. for no high mountain chains are present that are comparable with the Himalaya in Asia or the Cordillera in America. its effectiveness as a barrier will be apparent by comparison. Floating trees and roots afford a means of transport to many animals for an unplanned crossing of the destroy islands. In consequence the principal groups. does not present a barrier to flying equalizes temperature differences and nowhere forms a sharp climatic limit. a mountain range of considerable height not only forms a barrier to movement by its steep slopes. The starling.76 Barriers and Means of Dispersal than as a separation. for example. but body of water it animals. This is equally true of reptiles. There is no noteworthy difference in the mollusk fauna of the north and south banks of the Amazon. Small rivers and lakes are of no placed with the oceans as barriers may be of importance. The fauna of the northeastern strip is in . birds. and with them their fauna. In contrast.' 7 " The Mississippi at New Orleans is not a barrier to dis- which an increase in height of land of about 20 feet at the northern side of the coastal swamps divides two distinct sets of communities and limits the distribution of many species tribution. If the mountain range in question is older than an associated marine strait.

Barriers and Means of Distribution 77 the main like that of the Malay Peninsula and to a lesser degree like that of Borneo. their attendant climatic effects. The fauna of the Cape is also separated from that of central Africa by the Kalahari Desert. except for speaking. ness the effect of the tsetse fly carrying trypanosomes in restricting the of domestic ungulates in Africa. they extend from north to south. flying animals. they form veritable highways upon which polar or temperate forms may extend far toward or even into or through the tropics. Alps. they coincide in general with climatic limits and intensify them. Dromica. its cordillera extending from Canada to has been able to spread from Alaska to Patagonia. form more effective barriers to the distribution of snails than do straits. Carpathians. the endemic species of Sumatra are found south of the mountains. and the difference between these two parts of Sumatra exceeds the difference between its northeast slope and the Malay Peninsula. lines of dispersal for They thereby provide definite east-west or west-east many animals. . and faunae of diverse composition may then be brought into immediate contact. and contains a number of endemic genera. with Chile. on the contrary. Biotic barriers are also important even on a geographic scale. In Eurasia the principal mountains. and prevent crossing from north a sharply defined boundary for all terrestrial to south. In some. if sufficiently high. Manticora. such as the Himalayas. among the tiger beetles. especially when they are a nearly continuous series. the Pyrenees. If. practically range vanished from the drier parts of South Australia. Himalaya. 04 Mountains. extend from east to west. Deserts make animals. Tien Shan. The Sahara forms as distinguished from division between north Africa and central and south and only interchange occurs in the Nile Valley. as from Senegal to Mongolia. Hindu Kush. the puma Caucasus. This interchange is limited by the narrowness of the valley and does not affect the essential distinctiveness of the two faunae. the cold north is directly juxtaposed to the tropical south. Wallabies have. little a very occasional rock wallaby in the hills. wit- the faunul Africa.-is The fauna much closer to that of of Victoria south of the Great Dividing Range Tasmania than to the fauna north of the which agrees with that of New South Wales. as have all the interesting marsupials such as the jumping mice and the rabbit bandicoots or "bilbies. In America. The direction of mountain ranges has an important influence on range." The bilbies were apparently ousted from the country. and Altai. and Myrmecoptera. such as. When they parallel the lines of lati- tude.

Aus- .. far from land. which are otherwise poor in snails. a polar bear has been observed swimming in the open ocean 30 km. 54 effectiveness. These invaders now occupy the old and very useful they must have found the ready-made bilby warrens. 44 Twenty kilometers seems to be the maximum distance 9 that can be negotiated by the red deer. 5 km. they are said to have been taken by ships on frequent occasions. though a very few may adjust to brackish water. rang- ing to the Solomon and Fiji Islands. sea journey to the Keeling Islands. for the burrows. open country has a good deal of limestone in the soil either as rubble or as a solid sheet. Chile. usually arriving dead or dying. wide. they have occupied all retreat. The crocodiles are among the best swimmers. 30 km. from land. The speed of movement on land is relatively unimportant for the distribution of animals on account of the amount of time available. Reindeer also swim well and readily enter the water. relatively few terrestrial animals can swim any considerable distance. It has been found in the sea off the FinisIt is terre coast.78 actively Barriers and Means of Dispersal by rabbits. the Palk Strait. Europe that was covered by the ice sheet since Effective and active wandering by land snails is attested by the occurrence of the photopositive helicid snails on isolated areas and Cretaceous rocks in the North German Plain 'and by the relatively rich and uniform snail population of the scattered ruins in the Black Forest. 5 In the United States the colonies of the introduced Helix (Cepaea) Introduced European earthworms have hortensis expand rapidly. The European water snake Natrix natrix is a good swimmer.- thus somewhat curious that snakes tend to be absent or few in species on oceanic islands. for which the power of flight is of greater importance than swimming as a means of distribution.Many snakes also swim well. from the nearest resting place. has kept the tiger out of Ceylon. Amphibians are killed 51 by salt water. a distance of 30 km. fully hard labor. The means unequal of dispersal for terrestrial animals are very varied and of Active motion appears in the most complete gradation. The hippopotamus and polar bear seem to be the best swimmers among the terresjtrial mammals. Conversely. of Muschelkalk 1 displaced the native forms near the cities in California. two individuals of Crocodylns porosus are known to have withstood the 900-km. The hippopotamus swims the strait between the mainland and Zanzibar. Although earthworms and the area in northern its snails move slowly. which must make burrowing painand literally. Snakes have repeat11 edly been washed up on the Keelings. Leaving out of consideration the sea birds. but some survive the journey.

inhabiting at f annul provinces. Of the 6 species with complete wings. which is females. Many individuals make extended flights. and less of genera of limited distribution. However. is constantly maintained there by an influx from the south. moderate height. accessible islands The Flying animals are least limited in dispersal by physical factors. of the native oligochaete fauna. Thus 6 fully winged species of the genus Platycleis and 12 with vestigial wings occur in Austria and Hungary. air offers no barriers. Almost nothing is left. The arctic fox seen by Nansen more than 100 km. only the low temperature and lessened Deserts. and the the reindeer from ward. the morning-glory sphinx. such as ice masses. powers of rapid motion may enable animals to spread from the mainland to otherwise in- and from island to island. not one has both sexes flightThe mantid less. and seas of not too great extent are not barriers for good fliers. wolves have followed them. 43 Mountains and deserts form climatic barriers. forms with limited flight are usually restricted in The number this 8 safely be ascribed to their weak flight/' Flightsmaller than have ranges grasshoppers winged species. and the morning-glory sphinx ranges throughout the Old World. and only a few have weakly flying females. Thus the caribou have the Melville from Peninsula to Baffin Island and farther northspread the mainland to Nova Zembla. 7 Of 45 species of Orthoptera common to the desert and steppe faunae of central Asia and South Africa. north of Sannikow Land in the New Siberia Archipelago illustrates the distances to which swift runners may travel over the ice. the oleander sphinx. and Means of Distribution 79 and elsewhere. 2 range over all of Europe and 4 into the Mediterranean province. insects. there is great variation in the power of flight in different Among range. most two tera). When connections. are temporary. .Barriers tralia. groups of flying animals. mountains of density at high altitudes may affect flight. 5 are restricted to Dalmatia and one to the Swiss Jura. on the Antilles. and greater powers of motion do not necessarily enable animals to cross them. and have spread a considerable distance inland. is very large in the caddisflies (Trichop- may -' species ranging farthest to the east in the Polynesian Islands belong to the relatively small group without notable wing reduction in the The great power of flight of the hawk moths (Sphingidae) makes possible a wider distribution of their species than of other Lepidoptera: Celcrio cingulata has a vast distribution. 70 unable to gain a permanent foothold in East Prussia. while of the 12 flightless forms.

Mountaintops. Pantula flavescens sometimes appears in numbers on the Keeling Islands. without the formation of local races. from Ceykm. Several genera are worldwide in range or nearly so. and most species of fruit-eating flying by foxes of Malaysia and Polynesia are restricted to particular groups in North America. about halfway between Montevideo and the Cape of Good Hope. The large American noctuid Erebus odora has been taken on Tristan da Cunha. Some birds . and the common monarch butterfly. 1 Though flight reaches its highest development among birds. and is occasionally seen on ships not far from the European coast. and large flights of such forms have been observed. thus species of Myotis are found in every continent south of the tree limit in the northern hemisphere. which has spread across the Pacific via the South Sea islands to Australia since 1890 and is now also lish the most abundant butterfly at Sydney.- Bats. Among these are the thistle butterfly (Pyramcis cardtti). Various single species of Lasiurus and Lasionyctcris are extremely widespread Broad stretches of ocean do not scorn to be crossed accord.80 Barriers and Means of Dispersal whose pupa succumbs to the winter in central Europe. especially in the tropics. and dragonflies were observed daily above the water on a journey made from Singapore to Sydney during calm weather. 60 Some species of butterflies have a well-developed flocking instinct. The latter species has appeared at various places in India and Europe and on the Eng- and Spanish coasts/' A flight of more than' 20 came aboard the Novara 300 km. on all the The occasionally fish-eating Noctilio leporinus appears West Indian islands. may swarm with insects flying apparently carried up by the winds since they neither nor feed in that zone as adults. which is absent only in South America. Helena. also exhibit wide distributions of their species and genera. other species of bats have frequently developed special subspecies on the various islands of the Antilles. the death's-head has flown to St. 29 Great swarms of dragonflies frequently appear. bats of their own of islands. is caught from time to time in north Germany and even in Russia. and such specimens must have flown from points south of the Alps. ir> Papilla hector Dragonflies also include powerful fliers. in consequence of their powers of flight. Swarms have also been reported for the backswimmer (Notonecta glauca). They are especially there develop 4" There subject to being blown away from such exposed situations. there are great differences in the flying power of various groups. may be large-scale directed but quite unexplained flights of butterflies as described by Kingston in the Himalayas.

and these accordingly are the only kinds of spiders found in the Hawaiian Islands. Many in- of a goat chafer blown up to the glaciers in our high mountains. only three of these are common to both coasts. are poor fliers.water animals takes Storms place ways. In contrast with these. which many have relatively restricted ranges. but also small snails.. Victoria. The "Pampero. the stork journeys from East Prussia to Cape tundra and winter small areas. but actually becomes a source of food and a to rest. 17 Rains of caterpillars which were brought by storms are reported in south Russia. and leaves many not miles. two of which form separate races. and rains of springtails (Collemsects are in bolu} Balwyn. cited by . An analysis of the West Indian insect fauna by Darlington clearly indicates wind dispersal as Even tiny snails an important element in the origin of the fauna. such as petrels and gulls. and fresh. Spiders with their flight threads have blown onto rigging of ships 300 km. large larvae (Aegosoma scabricorne) fell to the ground during a thunderstorm in Basel. brings a veritable rain of insects to Buenos Aires and Montevideo. that birds are frequently transported by windSea birds from both the Atlantic and Pacific have been re- known corded in Wisconsin. It is well storms. such as Phisia gamma and Psiltira moiwclm. such as the pheasants. and thus transport only resting stages of Protozoa and small Metazoa. The other birds are restricted to relatively gallinaceous birds. the ocean not only is not a barrier. twigs. can be carried thus and may then give rise to a diverse snail fauna by 1:. spiders. butterflies. 21 radiative evolution. Colony and back every year. Powerful fliers of these groups are among the most place widely distributed birds." a southwest wind from the Pampas. The members of the heavy-flighted auks are so different on the two coasts of North America that. 17 The possibility of population of more remote islands by aerial transport is much better than was formerly realized.Finally. of 17 Pacific and 6 Atlantic species. have often been carried by the wind to Helgoland and even to England. and their Passive distribution of terrestrial in various eggs. insects. Only free-living spiders are transported by this means.Barriers and Means of Distribution 81 of prey such as the sea eagle and the barn owl have an almost worldwide distribution. the few cryptozoic Flights of continental spiders found there were introduced by man. carry dust. for some species. The accumulated records of accidental occurrence of European birds in North America north of Mexico. from land. Many shore birds nest in the Arctic in the temperate zone of the Southern hemisphere. myriapods.

Stream between latitude 39. 10 ls The woods bordering the La Plata have a fauna very different from that of the neighboring pampas. and even whole floating islands. A number down to the plains ' 7 another. include 44 species. and jaguars to the neighborhood of Buenos Aires. a floating in. it at once attracted attention It and was killed. such as the Amazon. that the actual establishment of a species of mammal or reptile in a new territory or island. few days. Congo. Ganges. and if this is taken into account the disproportion between the two lists becomes This clearly exhibits the influence of the prevailing westerly winds of the north Atlantic. rivers rivers. and the Indo-Chinese even large animals may be transported for long distances in this way. will die of starvation. have been observed at the mouths of many tropical rivers. American origin. amLarge phibians. 38 must be recognized.5 N. crocodiles. however. A boa constrictor was washed up on the beach of the island of St. mammals. of Alpine species of snails have been brought and have established themselves on the river banks: thus Cochlostoma septemspirale on the Danube at Kehlheim and Helix (Cepaea) silvatica from western Switzerland to Karlsruhe and Worms. have been recorded only from Greenland.Spix and Martins report a number of monkeys on a floating log in the Amazon. streams at flood time carry driftwood. a squirrel on another. could take place only under espeIf the sea journey lasts more than a cially fortunate circumstances. and a tiger cat and a huge caiman on animals. in consequence of this sort of transport. however.82 the Barriers and Means of Dispersal Check less No List of North American Birds (1931). and of 25 aquatic The British coast line is only about one-tenth the length of the North American.. Vincent coiled around the trunk of a cedar. was sighted repeatedly in the Gulf of 1000 extent about island sq. In the summer of 1892. especially snakes. list five "accidentals" from Europe Examination of the best com- parable of British birds 7t discloses records of 14 land birds of species. and thus transport not only many small forms but a few large animals. leaving for the North American continent. and especially small ones. Branches and trees and large rafts are carried out to sea by the and are then carried farther by currents and winds. The Paraguay occasionally brings large snakes. Masses of driftwood and rafts up to a length of 30 m. and brilliantly colored insects of northern origin. 2 than 11 of the 16 land birds in this list. and longitude 65-43 W. Flowing water often serves as a means of distribution of terrestrial still greater. tree trunks. .5 and 45.

the assumption of a large role for such rafting in the colonization of a given territory must be made with reservations. In to arboreal snails.' 52 Insects may be distributed in their pupal stages. This is an embarrassment of riches to account for the relatively few mammals found on truly oceanic islands.000 years (Knopf. a period ample for transport to great distances. 27 were still addition alive after 14 days. Barbour does not believe that rafting furnishes a feasible explanation of such distribution. This may explain the preponderance of snout beetles on St. with and without opercula. seem at first glance to be slight. Darwin and Aucapitaine have shown how resistant they are to sea water. In any case. attached to the bark or trunks of trees. " Matthew n and Barbour have critically examined the possibilities by rafting.Barriers reptiles and Means of Distribution 83 fast for longer periods. the number of probable successful Cenozoic colonizations of oceanic islands by mammals by this means amounts to 25.000. Employing Matthew's estimates of the probabilities of dispersal by rafts. The eggs of might come ashore unharmed in the roots of trees or in holes or crevices in their trunks. for rapid flotation. Snails. correcting the arithmetic of the original analysis. In general. pounded Even if the traveler has successfully reached the land. the strand-inhabiting Auriculacea are especially exposed to such transport. of colonization and allowing for the increased estimates of the duration of Cenozoic time. in Simpson 61 ). The geckoes lay hard-shelled eggs. unrelated except for the common adaptation . especially by mammals. especially wood-eating forms which pupate within the trunks of trees. colonization can take place only if both sexes of the species in question or at least a fertilized or pregnant female have completed the journey. which is would break over the floating object and endanger the travelers. of 100 snails kept under sea water in a perforated box. while Matthew thinks that it does. where they compose more than half the native beetle fauna. The ability to continue will even then new depend on the habitat conditions afforded by the Thus the chances for the colonization of new territory locality. in this way. calculated in 1947 as 75. among them 11 out of 12 species of Ericia. which require many months for developreptiles ment. Helena.000. the waves necessary might In a strong wind. and driftwood often is for days by the surf and only rarely carried directly ashore. The experiments of are good subjects for this form of transport. the chance nature of raft transport would result in a varied assemblage of animals. The chances are much better for forms with a dormant stage or with eggs that may be carried on these natural rafts.

States and the dyewood waifs to deer was introduced Europe There has been a great waif translike that of the banana waifs to the United Hamburg. and the to flies. for example. The wide distribution of pigs in the South Sea Islands is explained in this way. It is. and still more the feather and hair mites. (5T dispersal. as did cattle in Australia and pigs and goats on many islands. Partnla. as nonfortuitous. 64 Seafarers have often left domestic animals on isolated islands as a source of food for later visitors.84 for this Barriers and Means 5- of Dispersal means of dispersal. horses and cattle in the various parts of South America did so. The absence of other forms is then exactly in accord with the requirement of a "chance" assemblage by fortuitous ous species. beetles. it is highly unlikely that they arc any way parasitic. for example. fleas. are transported by their hosts. craneflies. commensalism under the term have been found attached and bugs. Primitive peoples caused this as well as civilized man. and then frequently became feral. and parasitic Diptera." in Pseudoscorpions. since there are 16 endemic genera of rodents with numer- The presence of 16 genera does not necessarily imply 16 since a group of related genera may evolve from a single landfalls. The majority of such . and goats on and established in central in Juan Fernandez. portation of different sorts. of course. The attachment to flying forms -4 dispersal of small non-parasitic terrestrial animals by temporary is of sufficiently common occurrence to be distinguished as a special mode of "phoresy. Pilsbry land snails of the Pacific islands as and Crampton n regard the spread by land connections rather common primitive characters of kidneys and genital organs of the genera TornateUina. entirely natural for parasitic forms to be carried in this way. The fallow similarly transported from the Mediterranean province. presence of the dingo in Australia is also attributed to the agency of man. The nest parasites of swallows and swifts. Gastrocopta. with the exception of the dingo. and others. and lice. on account of the placental fauna of Australia. The same is true of the minute external parasites of bees and ants. Game animals have been new localities. Some authors similarly regard the than by fortuitous dispersal. and it may be supposed that in some instances they had attached themselves in search of parasitic mites. Paul. successful emigration. Domestic animals were taken with him. By this duced on St. Intentional and unintentional transport by man has carried great numbers of animals to countries where they were originally absent. rabbits on Porto Santo means cattle were introand Kerguelen.

. 1 Gustav. 155. Results Voy. 1931. 9. 439). 18. 8). G. M." Zoo'/. Stomatopoda: 1-116. pis. 13:274-300.. "Land-Isopoden." Ber- gens Museums Arbok. Darlington. Bot. Verlaten Island. 526. 8:1-10. 14. Y. 418 figs." XI. 189). 1884. 10. Z. G. (p. Decken. being released from nomic the biotic pressure of their native land.). "Zur Systematik und geographischen Verbreitnng der Brachiopoden. 90:596-644. E. 1909. 599. 1904. Ostafrika. 16. Sd. 77:10-12.. 16 8. Rev. 1907. Acad. 1909. 1895. Bauer 12. 2 pis.. No. U. "Baron C. (p. (7) 14:403-410. tool. "Hjorten Norge (Cervus claphus atlanticus) . 1867. distribution. P." Nature. 1938.). Wash."' Ann. G. 5 figs. Jr. 2 figs. Buckle-Lund. A. but if they do. 6:1-31 (p. "The origin of the fauna of the Greater Antilles. 120.. 1869. 1909. 1911. ChalG. Lancaster. pt.. J. (p. 1916. R. The species inhabiting Tahiti/' Pub. they may develop into ecoThis whole subject of man's influence upon ecological pests. of the genus Partula. Dammermann. in the final chapter. zoogeography will be considered BIBLIOGRAPHY 1. and Sebesy. "Notes and observations on the distribution of the larvae of marine animals/' Ann. 2 figs. 2 pis. "The fauna of Krakatau. 4th ed. C. Gastropodenfauna von Basel. Stephan. 1898. "Unsere Land-und Susswassor-Mollusken.. p. Heligoland as an ornithological observatory: the result Edinburgh. Vcrhandl ZooL 16.. Univ.. maps (Abt. 228:1-313.. Ileinrich. Robert. Challenger. 185 ff. 1908.. Check-list of North American birds. Stuttgart. W. Set. H.. 13. Lutz: viii." Einfiihrung in die Molluskenfauna Deutschlands. (p.. 4. berg. Biol.. 22). Ges. 2 ed. 6. 4. Mag. 1922. Collett." Treubia. N. 54:175-263 (p. Brooks.: xix. 3:61-112. Rept. David. 1 pi. Bollinger. 7. "Some remarks upon Matthew's 'Climate and Evolution. Basel. "Zoologische Miscellen." Quart. 15. Allen. Carl Clans von der. 1 map. 27:1-10. Deutsche Exkursions-Mollusken Fauna. 18 pis.. O. Natural Hist. . Boulcngcr. 1916. 216 pp. Thomas.Bibliography waifs fail to 85 become established. 2. Clessin. 1909. 17:425-502. Douglas: x. Wien. 47). 627 ff. with discussion of dispersal of animals over water and through air. von der Deckens Reisen in /' Ostafrika in 1859-61Leipzig: 4 vols.. F. ZooL. Carnegie Inst.. (p. K." i Deutsch pi. American Ornithologists Union. "Studies on the variation. 13 figs." 5. Blochmann. Barbour. Gardiner. 34 pis.' Geyer. "Fishes of the Nile.MS. 11. lenger. Frauenfeld. W. 2.. "Report on the Stomatopoda collected by H. K. 3).. (p. 17. v. 126). 5 pis." Bull Museum Comp. Doctor's thesis. wiss. 3. Giitke. J. S. pis. and evolution Crampton. 1886. Niirn& Raspe: 658. A. of 50 years' experience. "Mammals of the West Indies..

(p. Barriers and Means of Dispersal Gudger." Natural History.. "Biological peculiarities of oceanic islands. 35. 344. ll:xviii." 30. N. Y." 2 pis. 400. Jones. Ges. 1912.. sp. Brehms Tierleben. Witherby: Proc. "Canard et anodontes. II..] London. 17:120. 24:171-318. 4fF. Sor. illus. 27:11-15. Gnlick. imt. "The fauna of Cocos-Keeling Atoll. N. Salvatore. 10:231-244. 40. "Kafer und Netzfliigler. Hesse. 2. 654. 1870:87-89.. Die Mollusken dcr Wilhelm. BioL. Theodore Monod. 2.. F. Zool. and G. 1867. 7:405-427. 24. 4 pis. illus. Hilzheimer. J. 1920. Wallis." Wiesbaden." Verhandl." Idem. 22:333-334. W. 1904. 37. Univ. Sri. "Climate and Evolution.86 19. J. Apple-ton. comestibles des cotes de France. Sta. Matthew. Max. 1938.. "Studien zur Zoogeographie. (4) 15:282-283. Tierbau und Tierleben." Mitt. C. Addison. 4th ed. 4:1367. G. 1-33 (reprinted.." 300. Proc. 1909." Treubia. 21:607-619. A geographical history of mammals. Lebedinsky. Kobelt. Acad. Johnston. Cambridge.. illus. ). xiii. Bull. 22. illus. 1928. Guerin-Ganivet. Y. 1870. (vol. 34ft. 2nd illus. Eduard von. R.). (p. Hartert. 503). J. 1896.. Acad. 131:1-12. 27. Neapel 19:51-3-763 (p. 36. Friedla'nder. Ernst. 41. l:xii. 530). ripede sur un poisson dulcaquicole europeen. Kuirea. 33. Lo Bianco. 4.. pub.. oceanog... 1909. Kew. and Franz Doflein. ed. E. llth ed. 5:363. naturforsch. W." Rev. 1935-1942. 157). Martens. figs. Richard. 21..." Nautilus. 29. Kolbe... "Die Preussische Expedition nach Ost-AsiVn." 31. 1901. fahigkeit des Batrachierlaiches gegen Austrocknung. B. H. London. xii. Aus den Wanderjahren eincs Naturforschers. 20. . A naturalist in Himalaya. Lyell. 223. 2 vols. Ibid. 1922.. "Die Saiigetiere von Alfred Brehm. Heinis.. 329. t 7 figs. C.. Monaco. Wood. 1909. (p. 1915.. ginalis.). Jena. 34. Soc. Quart. 369). figs." Deutsch Ostafrika. "Experimentellcs iiber die WiderstamlVerhandl. 1901. Klunzinger. pt. Berlin. Zoologisclier TeiL. Dinulesco. maps (p. London. 547). 1932. oceanogr. F. Zool." illus. sci. and Ludwig Heck. Mansion. maps (p.. 1897. 28. 82 figs." Rev. Lydekker. 1916. 1919. W. "Die Moosfauna des Krakatau. A. Kingston. "Presence d'nn c-ir32. 1877.. D. Zool. 1921. 1 pi. Hudson. 22:84. "Notes preliminaires sur les gisemrnts de mollnsques Bull." 615:1-32. [Notes on pampas birds. Zool. 1909:132-159. 1939. 1914. Y. "Ober einige Ergebnisse meiner Studien iiber die Rundkrabben des Roten Meeres. p. "Sphaerium corneum upon the tarsus of Dysticus marJ. Fischer: 2 vols. N." Ann. 26. 327). 39. 23. p.. "Rains of fishes and frogs. Menzel. Richard. Monaco. 1 pi. Cambridge 38. 2 vols. N. "Rains of fishes. supplementary note. Press: xii. W. and R. 33 figs. 3 text figs. W. (vol. 25. Conchol. 1-42. Sci. 1 map. 1888. Richard.. 1898. dcut. "Notizie biologiche riguardante specialmenta il periodo di maturita sessuale degli animali del golfo di Napoli. Basel 30:189-212. Principles of Geology-. palaearktischen Region. inst. "Ancyli adhering to water beetles. Kreiclel: viii." Ges..

" in idem. 58-61 (p.. 1903. 1867. pis. 1895. Notes by a naturalist on the Challenger. "Das Vorkommen von Sagartia luciue an der deutschen Kiiste. 2. Am. 184 figs.N. Sc/. R. being an account of various observations made during the voyage of 7/. 1908. 56. 4:1-114. Jahrb. 620. 334. the adventures of a x. (p. R.S.. P.C. 186. 1 map (p. 54. 27 figs. Albert. Saville-Kent. Barrier Reef of Australia. map. and Capt.A/. London.. N. R. 118). p. Pocock. 1921. N." C. 1879. 6 Ifg. 1S90. . Murray.. under the commands of Capt. 284. Stuttgart. "Reise in den ausserstcn Norden und Osten Siberiens." Zool Anz. 19 pis. 122). Macmillan: 47. Holt: 256. "The Echinoidea of the Swedish South Polar Expedition. Schmidt. II. 1919.. Bosmina benaehbartcn Ccbiete: I. 1871:175-186.). by atmospheric agencies. (p. S. 1 map (p.. Neu-Caledonien und die Loyalty -I nseln Reise-Erinnerungen eincs Naturforsclwrs. (p. 101 fF." 342)." Zoologica. illus.. 10 maps (p.. P/i/to. 59. (>(). 109ff. W. xviii. Francis. Flying fox and drifting sand. 3rd Intern. 1938.). 956). London. Y. (p.. Hist. 52:568-581.. 1900. 84 pis. lander: vi. 44. N.. illus. figs. ily 1903. 384). W. 45.. F." 48. Proc. Zool. Acad. F. 87 Mayr. Soc. Sir C. Sarasin. 46. "Monographic der Daphniden Deutschlands und der A. 2 1871. schwedischen siidpol. F. 49. figs.. 6:219-220. Exped. The Great 11. 58.).R. Y. E. Die Verbreitung der Oligochaeten.Bibliography 42.. Allen: 387.. J." Ergeb. Theodor. A.S. 53. "Concerning the development of frog tadpoles in sea Philippine J. von.. "On Mygalomorphae. Idem. Michaelsen. Ruhe. Congr. 1911. Turtles. pis. Berlin. Nares. 3 figs. biologist in Australia. Museum SI/A-/. Systematic* and the origin of species from the viewpoint N. crocodiles. Columbia Univ. "Allgemeine Biologic der Schmetterlinge. Pilsbry. Entomol. Pearse. 24). Monographic des Genus Bosmina. \atura1 39:38. 1912. in the years 1872-1876. Grossschmetterlinge der Erde. Miiller.. lizards. William. pt.N. S. Ratcliffe.B. 22 maps Bull. 401).. 51. Lehrman: 14 vols." Proc." Zool. 288 ff.. Ernst. the geographical distribution of spiders of the famSoc. map (p.. 50.. London. Adalbert. 52. Zool. 1910. A. coregoni im baltischen Seengebiete. "The general conditions of existence and distribution of marine organisms. Idem. McBride: 341. 1913. 1847-1875. A. R.. dispersal of non-migratory insects Trans. Moseley. "The genesis of mid-Pacific faunas. Y. 29 figs. Leydcn: 99-117 (p. Georg: x. pis. T. its products and potentialities. 2). Basel. ^ at. London. II.5-624. 55. 1843-1844": 7851094 (p. Middendorff. the "On water. (vol. 1903:340-368.. 109). 1913.. 5:281-343.. John. "Die Thierwelt Siberiens. Ferdinand. Friedof a zoologist. Congo 1909- 1915: Part I. 43. Pax. xvi. Heft 63:1-141. 7 pis. Mortensen. Press: xiv. Fritz. 1893.. The ocean-"." 7 Sci. Thomson. 7. 25. I. 12 pis. Challenger round the world.. K." 7-32. 1942. 57. 52:161-166. T. "Contributions to the herpetology of the Belgian based on the collection of the American Museum Congo Expedition. and chamaeleons.. K. 8 pis. Seit/.

71." Rept. and Richard II. 59:7081 (p. faunistische Studie. Basel 28. London. Ergeh. and general biology. letter to the editor. 154:138-142. G. Witherby. illus. Idem. 1943. The furred animals Sydney. Ellis. 60). iciss. 1942. 1942. 65. InsektenhioL. 64. Weber. 4 maps (p. 1892. A practical handbook of British birds. xxvii. S. 1 map. 92 figs.. 400. Fleming. 101). Taylor & Francis: 440.." Allgem. vols. "Der Rhein als Balm und als 2:28-65 (p. Die Tiefseefauna der Seen Mitteletiropas. 69.. 369). 1892. Reisc Ned. their physics." Proc.. Zschokke. Gcx. 126). 70. 27).. A. H. Y. Amphibia. 1947.88 61. 119-126 (p. 72. 8 pis.. 28 text figs. Witherby: 2 F. Schranke der Tierverbrcitung. Sweitz. 1 fig. naturforsch Ges. 1935. 1919-1924.. 68. 1907. Linncan Sot. 25 pis. W. pi. 21:480-483. ReptiJia I and (p. M. 76. Smith. 30. Vol. 77). "Die Tierwelt der Hochgebirgsseen. Idem. 4th Meeting Australasian Assoc. Max. Franz. 211-217 (p. Idem." PaleontoL. 246. London. 1919. 6:31. Z. Ges. wiss. 1:16-32. 182). 1917. 75. Eine geographiscliLeipzig. Sverclrup. Angus & 374. London. Zoo/. "A continental Tertiary time chart. "Bemerkungen iiber die geographische Verbreitimg der Mantodeen ( Fang-heuschrecken ) . Wicn. Georg. Troost.Australian archipelago. hot. 3:179-185... figs. Ulmer. Prentice-Hall: oceans. Adv. Neue Dcnkschr." Robertson: Z. 265 J. Garten. "Die Siisswasser Crustaceen des Indischen Archipels.. 1909. U. ." Zoo/. 2 maps (p." 73. 67. Verhandl. 1905.. "Uber die geographische Verbreitimg der Trichopteren. B. 63. 2:528-571. "Beitrag zur Kenntnis der Orthopteren Schlesiens." Verhandl.. 1087. Ibid. "Sauria. Friedrich. zool. nebst Bemerkungen iiber die Siisswasserfauna im Allgemeinen. Johnson. Martin W. 66. and Means of Dispersal /." 30:137-188. The x.. Insektenbiol. Barriers G. Klinkhardt: v. Ost-Indien. Zacher." The fauna of British xi. 37: vi. 68-80. The N. India. chemistry. 1900. of Australia. "On the fauna and zoological relationships of Tasmania. divisions indicated by the Vertebrata. Spencer. H.. Troughton. . map Idem. Simpson. pt. 7 charts.." 74. II. 02. 1865. 1911.: 82-124. F. "The biogeographic division of the Indo. "Die Tierwelt der Umgebung Basel nach neueren Forschungen. Werner.

however. A fairly marked climatic barrier is furnished by the 20-inch isohyet in the United States.e. the nature of these its may be very diverse. i. as is done by the mountains and ocean on the west coast of South America.. and these usually show a less definite boundary than do physiographic barriers.0. The Effect of Geographic IS Isolation THE AREA INHABITED BY A SPECIES sides IN GENERAL SURROUNDED ON ALL by barriers preventing thus limited in its range. has become very questionable with the growth of our knowledge of heredity. that "panmixia** would entirely nullify variation. in general. Thus arise subspecies. In the second place. Barriers of different kinds combine to produce an isolated area. species. and the species is As we have seen. with continuing isolalated related will. Climatic barriers also exist. which approximately coincides with the hundredth meridian. 89 . This makes possible independent differentiation of the two groups. since newly acquired characteristics will be restricted to that one in which they appear and will not beAfrica. The border of the range of a superior competitor or of an enemy or of a rich fauna may form a limiting barrier. they may separate closely related forms on the two sides of the barrier and thus prevent interbreeding. and tion. the presence of barriers operates to protect species within their isolated ranges from competition of rival forms or enemies that might be dangerous to their survival. the individuals within each group resemble each other more than they will members of the other group. thus enables protected forms to take tunities presented full advantage of all oppor- by their environment. come the general property of both. Barriers to dispersal have a twofold effect. In the first place. barriers further dispersal. The earlier belief that variations would be suppressed by crossing *vith invariant individuals. In two isogroups. or the desert and sea in North Cave animals are isolated by their negative reaction to light and by cave walls. and.

and . He even believed that this was the only way in which one species can become differentiated from another. in different directions in a relatively short and that the prevention of free crossing is an important element in selective breeding. even without intentional selection. from the evidence of distribution. The most important of the latter are. changes will individuals from another area can enter a partially isolated group. The experience of animal breeders shows that. if it removes a group of animals from enemies. ecological isolation.* 58 and. failing to take into account the other means of preventing interbreeding. and Alderney illustrates both of these principles. and competition. which similarly enables related forms by side in the same area though in different habitats or with different habits. that geographic isolation must have an impartant effect upon the transformation of species and the formation distribution of animals. Geographic the appearance of new adaptive modifications when the habitat conditions in the isolated area differ from those in the original home of the species. Isolation. Potent selective agents include climate. and by environmental selection of such favorable mutations when they do appear. '- who first concludes that even physiological and other differentiation arise geographic isolation. to exist side second. Guernsey. whereby the appearance of distinct groups within a species is possible without any geographic isolation. first. Wagner 8W of subspecies. "physiological isolation. isolated groups of domestic animals develop time. The development of the dairy breeds of cattle on the islands of Jersey. The varying effectiveness of barriers in separating interbreeding communities is of greatest importance to evolution. their effective mutational be brought with them. which occurs most rapidly when relatively isolated popu- lations are interconnected If 1 by emigration. Such mutational isolation particularly furthers change proceeds slowly. either large or small. but isolation may further the development of new races based on non-adaptive characters even though the habitat conditions do not differ essentially. in This matter has been reviewed by Mayr. may make possible the development of structures or colors that would otherwise be eliminated." brought about by a change in the bodily and instinctive characters connected with reproduction itself. The chief evidences of the importance of isolation in the origin of species are derived from the study of the geographic was the first to emphasize. food. soil.90 The Effect of Geographic Isolation There is both direct and indirect evidence of species change associated with geographic isolation. The transformation of a species may take place by germinal changes or mutations.

The tin-? Effect of Geographic Isolation of the 91 differentiation group from the aneestral form may be thereby retarded or hastened. The longer the isolation continues. nor but in an adjacent area separated by some barrier. not one is endemic. into wholly distinct species and even into new genera and families. however. The whole body of research on geographic variation. the red deer and wapiti. and for examples it is sufficient to refer to any systematic work dealing in a distant one. with longer periods of time. When. "' 1 7 These theoretic considerations on the in the role of geographic isolation evolution of species arc amply supported by the facts of animal distribution. the differentiation into distinct species is caused by geographic isolation. is This Jordan's with a large region. species. Physiologic isolation most frequently results from mutations of the generative organs of such a nature that two groups lose their mutual Infertility has thus come to be a criterion of specific disfertility. continues to confirm this rule. Rule* which seems to hold rather generally for expanding dominant groups of vertebrates but does not hold. such as Hartert's Birds of the Palacarctic Fauna. we do not find its nearest relative in the same district. of the animals that have colonized Krakatoa since 1883 from the neighboring islands. genus. this relation will tend to be reciprocal. There are certain exceptions in which intimately related forms occupy Tin 1 nst of the trnn "law" for such ecological generalizations does not seem advisable*. for ancient relict groups. The native fauna of the Hawaiian Islands. the larger is the number of mutations and the greater becomes the divergence from related organisms in The systematic value of these endemic forms subother regions. is largely composed of endemic species. The geographic races of the lion or of the zebra are completely fertile. and the European and American bisons. Thus. species. If we take a given species in a specified area as a starting point. family consequently makes possible a conclusion as to the duration and completeness of the isolation. Such new forms are confined to the isolated area in question. and then. With complete isolation. of course. tinctness. since it is in the highest degree improbable that identical mutations should become established in a species independently. by contrast. with numerous endemic genera and even families. which is now being carried out in detail for birds and mammals and some groups of insects. The isolated evolved form is endemic. and this is also true of such completely distinct forms as the various pheasants. mutual fertility need not be lost. the groups differentiate first into subspecies. .

in separate areas and have come together by emigra- if a sufficient degree of bodily or instinctive differentiation to prevent interbreeding has arisen during their separation. ranges through most of Eurasia and has entered North America from two different directions. 81 tively short time until the S. have They will then remain separate entered from three different directions since the glacial period. Among fishes. the gobiid Eviota and the blenniids Enneapterus and Salarias are represented by closely allied pairs of species in the coral reefs of Samoa. and have reached their present - common area of distribution only after postcharacteristically glacial migration. while Alaska from the Chuckchen Peninsula. is not surprising. and hence the more numerous the opportunities for independent variation. They have originated in three distinct areas from a preglacial common ancestor. is confined to springs . The animals of the Chicago area. Cambarus monongalensis. for example." The less the vagility of a species. The degree of .with clear water. south- western (Sonoran). the other. The Effect of Geographic Isolation 17 Thus Dunn regards the four species of Jamaican tree frogs of the genus Hyla as related stocks that have diverged into species sorted primarily by size at transformation. r>7 Closely related forms that now inhabit the same area may of course have originated tion. oe. the less it is able to overcome barriers. in view of the possibility of physiological isolaThat these cases must be cited as exceptions speaks for the high degfree of importance of geographic isolation. The powers of dispersal of an animal species may be summed up as its "vagility.92 the same area. Isolation is also to be seen in related forms that inhabit the same region but occur in different environments and have different habits or are sexually mature at different seasons of the year. diogcncs. Such differences This is illustrated by two closely reconstitute ecological isolation. One. lated crayfishes that occur together in southwestern Pennsylvania. and northern elements are distinguishable. a Eurasian thrush. 10 This tion. the more numerous are the areas that afford the condition of geographic isolation for it. C. oenanthe has reached a matter of only a relato It is two forms may be expected meet in Arctic America. Saxicola oenanthc lencorhoca reaches Labrador via Greenland. and Taylor writes of its grasshoppers: "This circumstance explains some cases in which two or three very closely allied forms inhabit the same area and occur in the same habitat. lives in marshes and other stagnant waters. The fauna of central Europe has a similarly diverse origin." The European wheatear. The same barrier may be of very different effectiveness for different species. southeastern.

unless they lead a planktonic existence by attaching to floating wood and pumice. howgreat. this is because of their plasticity of growth form. (9-10). there are no impassable barriers. neighboring geographic forms may be connected by intermediates. with the manifold conditions of their environment. especially aquatic snails. Widely distributed cirripedes tend to break up into local races. ever. a plasticity that obscures taxonomic characters. as are the subspecies of the wide-ranging Sicily. globularis (4-8). which by many authors as specifically distinct. and snails. while FIG. whose vagility is restricted to a brief larval period.The Effect of Geographic Isolation is 93 vagility of different groups of animals number of geographic races presented in inverse proportion to the by a given area. Thus species of birds are often represented in Germany each by a single subspecies. 13 The crested lark Galcrida cristata. and tticana Series of forms of Helix (Murella) scabriuscula (1-2). like most species of Lepas and Conchodenna. All the conditions are favorable to the formation of local forms among the reef corals. Transitional variation may be wanting are then regarded boundary between the ranges of vicarious forms. a strictly resident bird that rarely ranges far from its home. 6. and even neighboring reefs may be different. another with low vagility may be split into numerous local forms. When puma or those of the snail Murella (Fig. 6) in is The presence at the of such transitional forms the most usual criterion for subspecific classification. Occasional . vary almost from station to station. The gray sea eagle scarcely varies with a nearly world. from the mountains of Sicily. After Kobelt. whereas certain flightless ground beetles have nu- merous subspecies.wide range. their variability is very 14 in part. forms an unusually large number of local races. In the same area a species of high vagility may be represented by a single form. With a low degree of vagility mere distance becomes a sufficient barrier to prevent interbreeding. arranged in a geographical series from west to east.

became reconnected only with the and forest 1 '' 1 The corresponding species of a genus or the subspecies of a species represent each other in the corresponding environments of the two areas. it was pointed out to one of the authors (Schmidt) that the supposed isolating ridges are at most secondarily bare. The extreme fragmentation of the land snails of Hawaii and other South Sea Islands into species and subordinate forms has been ascribed to isolation of wooded valleys by bare treeless ridges. The two faunae then have a great similarity of composition. which would form a barrier to the dispersal In a conference with Messrs. These arboreal snails have speciated into a great number Zimof forms. In a search for other effective isolation factors.94 The Effect of Geographic Isolation hybridization. on the occasion of a visit to Honolulu in 1949. as in the crows Corvns corone and O. We believe that isolation by mere distance must have operated as an important factor in the speciation of these snails. Zimmerman. cornix. without occurring anywhere towithout being exactly identical. consisted in a continuous blanket of forest. and in Tahiti as well. whose continuous environment reduces the effectiveness of such barriers as exist. docs not appear to correspond to snbspecific intergradation. The same barrier may separate a whole group of forms from their relatives. or apparently so. gether. it was suggested that the lava flows of earlier eras of more active vulcanism may well have isolated numerous areas of forest of varying size. An excellent example of the effect of geographic isolation within a restricted area is supplied by the Achatinellidae of the Hawaiian Islands. reserving the term species tor the grade of differentiation at which reamalgamation with the nearest related form has become impossible. This is one of the reasons for the great absolute and relative preponderance of the terrestrial forms in number of species. and that the original condition in Oahu. merman 93 lists 42 species and 75 subspecies of the genus Achatinella from the island of Oahu alone." Such vicariation takes place on a large scale between the mammals of Eurasia and North America. Bryan. The importance of this distance effect even in so widespread and active a form as the leopard frog of eastern North America has been shown by Moore. We agree with Mayr in a much broader interpretation of the subspecies concept. apparently in association with topographic features. where . which of strictly arboreal animals. and Kondo. successional development of soil on the aging lava. 5 " Extreme geographic fragmentation is much more common among land and fresh-water animals than among marine animals. They "vicariate.

but teeth. The wombat takes the place of the marmots. Except for monotremes. and antelopes are represented on the two mountains by corresponding Such vicariation is demonstrable in eighteen species or subspecies/"' of which are included: forms. ptuifjutiiensis //. may include details of structure. The mammals of Kili- manjaro and Kenya exhibit a similar vicariation. the red deer and wapiti. tats are filled to continued dispersal. among pairs Kenya Kilimanjaro ('. Monkeys Lemurs Squirrels Cotttlmn iihyttxinirHx kikuycnxix a. and the European and Canada lynx replace each other. and the whole fauna in such an isolated region exhibits an intimate phylogenetic relationship that is wanting in areas that have been open ject to progressive extinction. The same phenomenon. ruudatux Lialago kikuyrnxi.Adaptive Radiation 95 the European and American beaver.1 /[cli(utriuriin Icrniac (S. the different habi- with animals of a single stock. but these are so transformed by adaptive radiation that they resemble the forms of diverse groups that fill corresponding niches in the environment elsewhere. and these occupy the most diverse habitats. the reindeer and caribou. the mammals are all marsupials. lemurs. mutational changes accumulate. the wisent and the American bison. a number of rodents. and animals of the same origin become so different that they live in wholly different environthe barrier is When ments. the civet cats. The Australian region affords an excellent example. and the dingo. that of the wolves. the European elk and the American moose. Thus the different habitats come to be filled by related ani- mals. monkeys. such as the form of the and is sometimes so great that one may speak of convergence. the dasyures. flying phalangers replace the fly- ing squirrels. elephants. squirrels. The bird faunae of the north and south islands of New Zealand exhibit the same relation. shrews. Adaptive radiation an old one and the isolation has been long continued. . hyraces. inate species" is referred to by Jordan under the term "gemconspicuous on the two coasts of Central America (see Chapter 7). and the kangaroos fill the place of the larger herbivores. the Tasmanian wolf. various mice. umlulatus ralidti Hy nires Prurtirhi rnnrnhni/i P. with their fauna accordingly subIn the isolated area. river hogs. The resemblance is not only one of habits and in part of outward ap- pearance.

the concept of ecological equivalence. from the herbivorous single family forms to the predatory ones. is so great that direct relationship between the two forms. in it. related to their common adaptations for burrowing. whitefish. the banks of mountain streams. or worms and snails. and in the length and form of the tail. garded in this sense as ecological equivalents of the horses. These are especially remarkable in paralleling other distinct genera of the oceanic Cumacea. which is rare in the ocean. and small Synallaxis are found in the most barren parts of Patagonia. Fifty-five 33 genera of this family are now recognized. and this has representatives of all types. Among the South American birds the ovcnbird family (Furnariidae) shows a similar adaptive radiation. one-third of all and catfish. or the pampas. as the moles Such forms as the plains-dwelling kangaroos may be rejust cited. the dovekie ( Alle alle). The characins include about the fresh-water fishes of South America. with hundreds of species. or insects. burrow trout. whereas in the Caspian it has 13 very diverse species. they inhabit the rainand hot dry valleys. they are correspondingly varied in appearance. in length of limb and toes. replacing birds of the most diverse families in other parts of the world. Bodies of water isolated for long periods may acquire a similarly radiating fauna. the southern. somewhat less in general appearance. in the form and size of bill. among other examples: Pseudocuma diasty- . live in bushes. undergone similar adaptive radiation on the 46 Islands. pike. and differ in size. corresponding to our carp.96 The Effect of Geographic Isolation Thus the resemblance between the mole-like marsupial (Noton/ctes) of Australia and the placental golden mole (Chrysochloris) of Africa. though without morphological resemblance. forests climb like woodpeckers. they feed on seeds. thus. Galapagos The auks and penguins are analogous groups in the arctic and antarctic seas. They resemble each other greatly in habits. they reach the high mountains. may properly be extended to include the presence of taxonomically unrelated ani- mals in remote parts of the world but in similar habitats. 17 Adaptive radiation frequently produces ecological equivalents. Closely allied vicarious forms are likely to inhabit the same ecological Cope assumed a niche. the emperor and adelie penguins and their relatives. or Darwin's finches. which belong to as many different families. however. members of the sparrow family have (Fringillidae). they live on the ground. The extinct great auk. 14 species in all. The crustacean suborder Cumacea is represented in the Caspian Sea by the single genus Pscudocuma. and razorbilled auk (Alca torda) are the northern forms.

often very small. while those that stricted to special areas. lopsis deformis. which are especially numerous in coastal waters. such as oceanic depths. 8 . abbreviata. This applies particularly to forms with slight vagility. Geographic isolation in the ocean The degrees of isolation in the ocean vary in its different parts. 11 The conditions are different with the inhabitants of shallow water. 3 are endemic. In the tropics the Indian and Pacific faunae agree closely. as well as the temperature factors. Thus the Gulf Stream forms a temperature barrier that divides the species of craspedote medusae into two groups.Geographic Isolation hides resembles the genus Diastylis. whereas the Atlantic differs from both. but only 2# are Indian and 5? IndoN Ascidians are represented on the east and west coasts of Pacific/' northern North America by parallel series. P. and their distribution is dependent on the nature of their larval stages. freshening of the water at river mouths. peninsulas. to the pelagic forms of the Pacific that are absent in the warmer parts of the Atlantic are for the most part merely replaced by allied forms. 63 are endemic. and the number of species in each genus is larger. and the fresh water . Those with ciliate larvae are widely distributed. the genus Campylaspte. Of 52 actinian species on the east African coast. The sessile actinians. whose effectiveness depends upon the eurythermal or stenothennal character of the animals concerned. but within these the only surface barrier on a geographic scale is that of temperature. in the Ocean 97 Eudorel- P. frequently form endemic species. Connected marine coasts form highways of distribution for such forms. The ranges of species are consequently smaller. the actively swimming pelagic species tend be world-wide in their distribution. all the actinians of the New Zealand coast are endemic. nature of the bottom.- Canary Islands. Their distribution is limited by more numerous barriers. in have no free-swimming stage are rewhich they may be extremely abundant. The continents separate it into several major divisions. 21% occur also in the Red Sea. Geographic species and subspecies that replace each other in adjoining areas are frequently developed. for ex- ample.Generhowever. The construction of ship canals at Suez ( 1869 32 ) Of the 10 actinians on the coasts of the and Panama for ( 1914 ) marine organisms has not opened free channels of communication across these isthmuses to the extent that might be expected on first thought. The open ocean has few barriers. the arctic campy laspoides. whose members correspond exactly but are distinguishable specifically. The Panama Canal is not at sea level. ally speaking.

the specimens from the . may afford sufficient isolation for the divergence of species. like Ticrra del 15 ' Fuego. Suez Canal in- clude: (a) Plankton-bearing currents are deflected from the Mediterranean entrance at Port Said. the The salinity of whose waters varies enormously from hour to hour. has not as yet been carefully determined. is at sea level. the bottom mud is devoid of life. In the flounder Plenronectes flesm. (b) There is oil conBitter (c) The canal passes through the Lakes with heightened salt content. brackThe Suez Canal ish. Helena. In Great Bitter Lake. fishes of St. The factors that tend to prevent ready passage of the however. 40 different. It is a member of one of a small group of families found nearby inhabiting salt. Littoral sharks waters leads to specific transformation in and rays are scarce about New Zealand. despite these " A ctenophore obstacles. The high salinity of these lakes probably acts as a barrier even though bryozoans and isopods are carried through their waters on the bottoms of canal barges. the Falkland Georgia. this lies directly over an ancient salt bed which is gradually being redissolved and presumably will vanish by the end of the present century. a thoroughfare for marine organisms. during the several The tarpon (Tarpon atlanlicus) has to ships completed transit from the Atlantic to the Pacific. appear as vicarious subspecies.98 of The Gatun Lake kills Effect of Geographic Isolation most animals attached its hours required for passage. 30 Restriction to coastal the fishes also. while those of the Hawaiian Islands are mostly specifically.1 was recorded from the Palestinian coast in 1942. It resembles those found in the Red Sea. for sessile or mainly sessile animals. South Shetland. lack of vagility. though not generically. and for ten months of the year differ- ences in water level and the prevailing winds cause currents in the canal to flow northward from the Bitter Lakes. and this one may well have come through the Suez Canal. 11 ' Out of 65 of the marine but form a number of endemic species. which still Even mere possesses a considerable degree of freedom. South which the same* types Islands. and fresh water more or less indiscriminately. Ctenophora were supposedly absent from the Mediterranean. 17 are confined to the island littoral/' The coastal fishes of Micronesia and Polynesia are closely allied. extent to which animals pass through the Suez Canal. (d) A tamination at the Red Sea entrance. The littoral fishes of the Antarctic Ocean form groups in of species in definite areas. it is not. and Grahamsland. without any special barriers. further obstacle to free passage is presented by Timsah Lake. evidently on account of the isolation of the latter group.

Pacific. and Indian oceans. the "left-sided" forms are only 5.Geographic Isolation in Inland Waters 99 English coast (at Plymouth) run 30$ "left-sided. in extent. the 13-armed on a stretch of the Greenland coast. water are accordingly extremely favorable. almost uniform number In predominates. their opportunities for dispersal being for the most part confined to a short larval period. 13 agree also in starfishes of the genus Pentacer aster. or a pool is surrounded is by water. The same phenomenon is true for great variation that each individual branch may be subject to special 14 conditions. the dissolved substances. less than 30 exceed 1000 sq. at others the 11 -armed. present a great variety of form. ranging through the enIndian and Pacific oceans. and their environmental conditions have such conservative standpoint they latus. the some places the 10-armed form 12-armed at Tenby on the English coast." whereas on the coast opposite. The great majority of inland lakes an as island land by are small. A very small. a widespread genus in the Atlantic. Rivers are separated by land at their sources. equally effective barriers for their fresh-water inhabitThe environmental conditions. organic and inorganic. in sharp contrast with the uniformity The theoretic conditions for the transformation of species in the sea. have been united into a single protean species by the demonstration of transitional forms. 1 -' all belong to a single species. by the sea at their mouths. P. by either active . and there are almost as many modifications as branches. inland swamp. The variability of the wide-ranging inhabitants of fresh water is great. and many are lake. such as the development of the dorsal skeleton and number of paxillae. the kind of bottom. 87 Reef corals are famous for their diversification and adaptability. !H In the starfish Solastcr pappostis of the northern seas the number of arms German varies from 10 to 14. ants. especially in standing waters. Thus 32 supposed species of Spirastrella. isolation of small areas especially characteristic of pond. km. the specimens from given areas other characters. but is conin fresh trolled in part by the transgression of barriers at times or passive transport. but so many transitional and intermediate types are present that from a tire The mamisome sponges whose vagility is slight. while those of the Kattegat are 14-armed. the vegetation of the shores. but specimens from a given locality have an of arms. all combine to give an individual character to almost every body of water. are extremely variable: the daily and seasonal temperature range. Geographic isolation in inland waters is Geographic inland waters.

e. thus the species L. 7 Haas reduces the hundreds of named forms of palearctic unionids to 20 !> FIG. Anodonta. and in the coloration of their outer and inner surfaces. Many smaller lakes of that region also have their own species or subspecies. lacmtrina. var.100 The Effect of Geographic Isolation Examples of the multiplicity of variations in fresh-water animals are striking. d. the appropriately snails of the families named Dreisscna polymorpha. Outline. var. but they are completely united by intermediate specimens. The lakes of the glaciated area in Europe are inhabited by fishes of the salmonid genus Coregonus. var. The species of cladocerans. thick shell) . species. Lakes Huron and Michigan share three such forms that are not found elsewhere. Each of the Great Lakes of North America except Lake Huron has its own form or forms of one or more species of lake herring or cisco. hartletti is known only from Siskowit Lake on Isle Royale in Lake Superior. forma typica. piscinalis. The genus Pisidium. size. while French students later recognized more than 200 for the same area. 7). vary in form from lake to lake 69 (Fig. thickness of shell (Fig. var. The forms differ in outline. After Buchner. especially of the genus Bosinina. 8). anatina. was divided into 26 species by Kuster. according to Clessin. 7. cellensis. whose variability is such that each well-isolated lake is inhabited by well-defined subspecies of one or more forms. salmonids of the genus Leucichthys. Thus the genus of mussels. represented by 2 species in middle Europe. c. the degree of divergence being roughly proportional to the distance between the lakes. and the Lymnaeidae and Planor- bidae exhibit a similar variation. and shell thickness (heavy contour of five forms of Anodonta cygnca at their centers of distribution: b. Variation in the environmental conditions is in general less exten- . average size.

Summer Lake forms of Rosmina corcgoni from various Baltic lakes: H/. with their high content of dissolved humus materials. The bog-fed brooks of Scandinavia lack the alpine tur- faunae concerned in rivers of bellarians. a species for each river system. Lake Stein- Paarstein. krug. Lake Lake Dlusitsch. 6. Lake Wol/ig. though some important variation in the composition of the water of brooks and rivers does occur and affects the the same way as in lakes. The black-water South America. Fie. 39 In North Plata.nnin. 5. The by semi-arid mollusks. and Celebes."- plains. After Riihe. is inhabited by different species of Barfct/s. differ in the composition of their faunae from the neighboring streams. 8. 7. 28 and in the Rocky Mountains each river system tends to have its own species or subspecies of trout. 2. isolated rivers of Patagonia. Africa each river . The fish genus Rhamdia (Pimelodus) has split into at least The widespread south of the La 15 species in the rivers of Central and South America. flowing into the Atlantic and separated in the rivers than in lakes. found in Africa. Bengal. has 8 different subspecies besides the one in the Nile. 3.Geographic Isolation in Inland Waters 101 sive in flowing water. 1. Lake Luggewiese. are inhabited by distinct faunae of fresh-water fresh-water crustacean Caridina niiotica. Lake Wolzig. These are exceptions. 4. The effect of isolation in preventing interbreeding is more marked different large drainage basins of 13 The wellGermany have different species and varieties of mussels..

which 1 The fresh-water sponge Ephydatia them against the drying up of their habitats. with its highly speprotect cialized fauna of eleven genera of fresh-water mollusks. Australia. also have a wide distribution. adapted for wide distribution by their resting stages. bidae.contained only 3 new forms. The animal species of fresh-water plankton are to a large extent cosmopolitan (copepods. Ancylidae. Even the snails and mussels of inland waters have surprisingly large generic and even specific ranges. to turbellarians of fresh waters are very widespread. Even the exceptionally isolated New Zealand. There is thus a contradiction between the high degree of isolation of bodies of fresh water faunae. the great number of species that might be expected from the abundance of isolated areas is not realized in fact. such as Ranatra and Notonecta. Among species. and cladocerans ). crustaceans. insects. A satisfactory Belt. has only one endemic genus. and are not genetically permanent species characters. but with the exception of the Centropagidae. These are all.-"-' Of bottom-dwelling forms. though they are incapable of active The families Lymnaeidae. All the fresh-water coelenterates and sponges of Australia belong to European genera. fresh-water animals have a world-wide distribution. Planormigration across land barriers. America. and Unionidae have a world-wide range. and of 213 rotifers from the same territory only 21 are endemic. and Cybi. most sponge genera have a very wide range. 10 fluviatilis occurs in Europe. Protozoa and rotifers are very widespread.102 The Effect of Geographic Isolation In spite of the vastly increased opportunities for isolation in fresh water. and South Africa. bugs.ster 14 Some genera of water tripunctatus has a range nearly as extensive. Thirty-six species and varieties of rhi/opods from the region of Lake Tanganyika 7 . Many invertebrate of mollusks. Two species of Hydra and three of Spongilla are regarded as specifils Many of the rhabdocoel cally identical with the European forms. for example. The modifications and fishes cited above are in part transitory. arising from the direct influence of the given external conditions." be sure. North and in the Malay Archipelago. the dytiscid Eretes sticticus is present on five continents. Physidae. Thomas who explanation of this contradiction was presented by called attention to the transitory nature of lakes . The presence of the and the low degree same species of differentiation of their in the tropics and in the temperate zones is not explainable by a greater uniformity of environment in fresh waters as compared with terrestrial environments. The same phenomenon characterizes fresh-water The are more tropical and temperate representatives of the Dytiscidae alike than in any other family of beetles.

000. Many Siberian lakes are in process of drying up. Lake Constance is constantly being filled by the Rhine. The known to geologists as surface of the latter at its that of the present Pyramid Lake.4 was a real lake only 50 years ago.. 160 sq. a number of large fresh-water basins in which the faunae differ strikingly from the usual fresh-water aspect and exhibit the results of long-continued isolation. km. above Lake Ngami in Bechuanaland now a marsh in process of dis- a longer diameter of more than 100 km. however. km. They have numerous endemic species and genera. at 50 N.. plant remains raise the level of the bottom. Lahontan.Geographic Isolation in Inland Waters 103 and streams in terms of geologic time/"' Ponds and lakes in general have a short life. including the well-known phenomenon of stream piracy. for example.000 cu. a number of factors combine to destroy them. Then are.m.. the Sary Kupu. All these basins are characterized by large size and considerable depth. their faunae must be extinguished. and their genera are strongly differentiisolated 1 ated into species adapted to the various habitats available. at least such elements as were effectively and hence on the way toward changing to new species. desert valleys of the Great Basin in North America were lake basins Many lakes of vast extent. and. 440 m.500 years. 00 With the disappearance of such waters. deep. in . was later broken up into 20 considerable lakes. and Matano 800 sq. km. 420 sq. as they dry up. is reckoned at 45. Shallower ponds and oxbows. have also operated to break down the isolation of their faunae and to further the general phenomenon of wide distribution in fresh-water animals. gradually disappear as the result of the encroachment of the vegetation. whose banks are covered with vegetation. The the principal ones are Lake Baikal (35.. 590 m.000 years. Rivers and brooks fill the lakes in mountainous regions with detritus. deep. are their faunae During a part of the Quaternary. 203 m. marsh and bog plants grow out farther and farther toward the middle and limit the open water until it is The process of extinction proceeds rapidly in finally entirely gone. the destroyed. The frequent intercommunications between river systems. deep). and it is estimated that this process will be completed in 12. into which the Rhone brings an annual sediment of 2. 1706 m. and that of Lake Lucerne is placed at 23.. latitude. three central lakes of Celebes (Posso.. km..000 sq.000 years. Switzerland. a senescent lake when once a bog margin and false bottom have been established. it is appearing." 8 and rivers fall victims to drought. formerly an elliptic basin with Lake Bonneville and Lake maximum was 167 m. the life of Lake Geneva. Towoeti.

km. Geographic isolation on land Islands arc unquestionably the most effectively isolated of land areas. and depths of 1435 in.." (See Chap" 11 ' ter 18.) In faunal composition the Caspian Sea. compares more closely with these bodies of fresh water than to any of the partially isolated seas.000 sq. d. more Endemism on islands is the difficulty of reaching the island most frequent in forms for which is most extreme. proportioned to The amount of of endemism is inversely accessibility thickly populated areas. Tanganyika and Nyasa. Paramcluniti damoni. The strength and regularity and direction of the wind are also factors in populating islands.000 FIG. c. so that new in- . with a surface of 439. Lavi- geria diademata. km. 4 Recent studies of the fresh-water fauna of Lake Ochrida in the Balkan Peninsula show that it is to be added to this 80 list.. depth). Tiphobia horei. They are comparable with inland lakes and ponds in being surrounded completely by an effective barrier. Islands therefore present an abundance of endemic species and genera. probably with some other Balkan fresh waters. 9. but they differ from these bodies of water in their relative permanence. Tanganyika snails: a. Limnotrochus kirki. sq. each with an area of about 35. /. usually the nearest mainland. and a depth of 946 m. After Luutcrhorn. respectively. and even some endemic families and orders. and 800 m.104 The Effect of Geographic Isolation 45 and the Great Rift valley lakes. the the island from other.

The degree of isolation in either case may be the same. The Hawaiian Islands are the most completely isolated archipelago. The family Achatinellidae is especially notable. Carclia to Kauai and Tluianwnia Oahu. The amount of endemism on an by the mode of its Endemic forms consist of large series of species of certain genera and smaller series of related genera. which combine into families. with 12 genera and subgenera and more and mantle/ than 300 species. Unfortunately this very amount of differentiation is usually the only evidence available for the estimation of the age of an island. the eggs of the fresh-water snails are easily dis- tributed by aquatic birds. Similarly. however a phenomenon to be examined in more detail later (Chapter 26). They form a group of volcanic islands extending from northwest to southeast for about 475 km. as in the Polynesian islands. Speciation in these flying forms may be explained as the result of isolation made effective 74 by the development of homing instincts. in Melanesia fruit bats with powerful flight have formed numerous endemic species in several archipelagoes. In the very recently repopulated Krakatoa no differentiation has taken place. distant from the American coast and as far from Samoa. The composition of the fauna is different. by the breaking up of a former continental connection or by independent development as a volcanic or coral island. but their land snail faunae for the most part differ from island to island. where there are endemic species of birds although the lizards are identical from island to island. is confined to the Hawaiian archipelago. and more numerous in the latter group than among bats in and birds. Pcrdicclla and Newcombui are found only on Molokai and . whereas the land snails' eggs are dependent upon driftwood and storms. On very completely isolated ancient islands the genera are frequently split island may or may not be influenced whether origin. Some 51 to genera are confined to special islands. Endemic species are thus sometimes more numerous among mollusks and reptiles than among insects. snails are endemic.Geographic Isolation on Land 105 crements of the parent form are unlikely to follow. The duration of the isolation is naturally of great importance for the amount of change undergone by the fauna of an island.. The Antilles have a large fauna of fresh-water snails common. The converse relation may appear. All the species and three-fourths of the genera of land well separated. with only a general relationship to the widespread Polynesian genus Partula and with highly primitive characters in its sexual apparatus This family. and the individual islands are up into numerous species. more than 3000 km.

and only 5 out of 24 species are Certain finch-like forms that look like thrushes are wholly confined to the islands. from which these islands take 1 name. Each island containing a bit of the original forest had Hcmignathus tends its special species of each of the 9 genera. species South America. ljy The aquatic birds belong for the most part endemic. the mocking bird Ncsoininins. 9 are endemic. the flightless cormorant Nannopterwn and the fork-tailed gull Creagrafi. barn swallow. and 7 land birds. and osprey. whose dispersal may have been favored little wind. Sixty-two of the 108 species recorded are land birds. Of the 59 genera of birds.106 The Effect of Geographic Isolation Maui. Wasps of the genus Odijncrus."' The genus of lizards Tropidurus is represented by separate species or subspecies on each of 12 islands. one from the coast of Peru and one from the gulf of Panama.. most of the 76 named oceanic currents. west of Ecuador. and 9 genera with about 40 species form a special family. Important elements of the fauna are not confined to the Islands. in contrast to earlier reports in Fauna Hawaiientains is ' IL> sis of about 2700 endemic species out of a recorded total of 3325. exhibit a this greater development in Hawaii than in any other area inhabited by almost cosmopolitan genus. by 12 but the genera are without exception found are endemic. Each of the valleys radiating from the moun- Zimmeroften characterized by a special series of species. The endemic except for the bobolink."" . The giant land tortoises. their 5 1 See citations by /iimiuTiiiaii. with more than 100 species.7 man in 1948 knew of over 5000 species of Hawaiian insects. to widespread species.*'" The Azores present a contrast with the above-named archipelagoes. archipelago two ocean currents flow past them. the Galapagos ground dove Nesopelia. to and have 2 species. one large and one small. Some of of land birds genera these species occur on only one island. are represented by 14 species on the 9 largest islands. on This each of the larger islands. These include two water birds." The Galapagos is Islands present similar faunal characteristics. are little specialized. of which 3722 were endemic. the Drepanididae. and also in Central or 5 genera of the endemic family of finches Geospiziclae. of include 49 the 59 endemic forms. and Anutstra and Leptachatina have endemic species on each of the principal islands. sometimes Their relations are primarily with the formation of subspecies/ American. situated on the equator about 900 km. others on several. and all these are endemic. but they are in a region of relatively The land snails.

3 distinct families have been developed ( or preserved ) These are the primitive insectivores Centetidae. from the Portuguese coast. This group composed of 9 volcanic islands 1400 km. of amphibians 40$. The rich amphibian fauna and the diversity of the fauna of lizards and snakes indicate connection with Africa. of it necessary to place The dominant genus earthworms is Among snails the Kt/notus. the rodent family Nesomyidae. are represented almost entirely by peculiar genera. but 3 are endemic.Geographic Isolation on Land as their fauna exhibits 107 is much less endeinism. Matthew/"' to be sure. Lemurinae. . either identical with or very allied to the American genera Constrictor and Boa. of land snails 79$. The dependence of the formation of species under isolation upon the vagility of the groups of animals concerned is well illustrated by the fauna of Celebes. and to 1 may be assumed the resultant peculiarity of its animal life makes the date of connection far back in geologic time. with 7 genera and 23 species. with 12 genera and about 50 species. with few elsewhere. exclusive of bats. and still others burrow like gophers. with 10 species. Of the species of birds 28% are endemic. The Madagascan fauna appears to have had a long and undisturbed period of development in isolation. while Ampclita and HeHcophanta Forty-two of the 46 Madagascar genera in the cetonid The amphibians and reptiles family of beetles are confined there. Through the long-continued isolation. the family otherwise Fiji Islands. less than 30 breed on the islands. genus Tropidophora has a great number of Mada- are endemic. of mammals 40$. confined to Madagascar. occur in all Madagascar. with 7 genera and at least 12 species. have become differentiated races. and of land planarians 91$. was once united with large enough Africa. of . Islands close to the mainland are often little differentiated in their reptiles 36$. closely found only in the Americas and in the and two of boid snakes. gascan species. The Centetidae have undergone adaptive radiation. some quite without existing related forms in Africa. some live like shrews or hedgehogs. others in the water like muskrats. regards Madagascar as an oceanic island never connected with the adjacent continent. an island almost be called a small continent. Of 28 genera of mammals. Two endemic genera of iguanid li/ards. It sufficiently to and only 5 of these be recognized as Azorean with certainty that Madagascar. Of the 122 kinds of birds recorded. and the subfamily of lemurs. The only li/ard is identical with a Madeiran species.

high plateau of Tibet. 77 It is true that about one-third of the 41 fresh-water fishes are endemic. of 65 mammals and 64 reptiles very few are confined to it. bounded on the north by the Kwen-lun. a series of small islands that of the east coast of Sumatra. in spite of the great uniformity of the insular conditions. whereas all the similar deer on Sumatra and on the Malay Peninsula north to Tenasserim belong to a single species. though with 2 endemic genera. even so. Thus the genus Coercha of the family Coerebidae occurs in Central and South America and in the West Indies. The Tibetan mammal fauna is perhaps the most peculiar of any on a continental area. off In the Rhio-Linga Archipelago. island Mediterranean island groups. less 4 There can be no question that the habitat condiTragulus napuJ' tions on these small islands are less varied than in Sumatra and the Sumatra. disregarding the Tas- manian wolf. there are 19 in the Antilles. with an area of about 34 % than 8 well-distinguished subspecies of dwarf deer. there are no mainland area. but isolation operates endemic. which are recently extinct in Australia. An apparently innate variability has led to extreme fragmentation of the lizards of the genus Laccrta into recognizable races. Sarcophilus. the species are identical or in vicarious pairs. the genera and often the species are identical with those of the mainland. The islands of the Aegean Sea each have special varieties of the clausilid snail Albinaria coerulea. and Nan-shan and on the south by the Himalayas.108 The Effect of Geographic Isolation fauna. they are mostly local races of widespread forms. Environmental difference in the isolated area is also an important factor for the transformation of isolated species. or of fresh-water fishes or land and fresh-water snails. The vagility of the respective groups governs the amount of endemism in species. 41 lation as by 53 island and islet by islet. Thylacinus. and the Tasmanian devil. 5 of the 28 genera and 30 of the 46 species are endemic." . with an endemic form on each to of birds island. and only 13 out of 63 land snails are endemic. A few areas are surrounded by mighty barriers. there are no endemic genera of vertebrates. Tragulidae. such as the 3600-m. only 10% of the species of birds are ' 71 compared with 81% of the land snails. 18 The fauna of Tasmania is closely similar to that of Victoria south of the Dividing Range. Trinidad is zoogeographically indistinguishable from adjacent South America. Altyn-dag. with 13 subspecies on the mainland. in the - The mainland does not afford the possibility of such complete isodo islands. as produce new forms without differences in habitat.

and the Himalayas. snails in different valleys in Hawaii furnishes a still better illustration of this point. As they are photonega- they are unable to migrate from one hollow to another unless there are subterranean connections. Mountain faunae accordingly afford numerous evidences of the formation of species and subspecies The mountain ranges that extend in interresulting from isolation.* The mountains of middle and south Germany have numerous geographic races of the carabid beetle Carabns sihcstris. and are probably confined to this mountain. series from the rupted Pyrenees to the Himalayas and toward Syria and Abyssinia have their special subspecies of ibex. nubiana in Sinai. The tive isolation of cave animals air.Geographic Isolation on Land 109 The upper levels of high mountains or of mountain ranges that rise from plains to a considerable height are as effectively isolated as islands for stenothermal cold-tolerant animals for which the warmer lowlands are impassable barriers. one of the 4 species of fishes belonged to a new genus and species. of 21 mammals 6 were new. . C. C. The photonegative birds of the forest floor in the Amazon basin are effectively isolated by the broad rivers and their overflow areas. sibirica and its varieties in Persia. C. and the beetles of the genus Carabus and many Lepidoptera form special variThe well-known isolation of eties in the isolated valleys of the Alps. of 161 birds of 128 genera 41 species and 6 genera were new. C. how- Capra pyrenaica in the Pyrenees. severtzowi and C. their habitats are and avoid dry as isolated as ponds for fishes or as islands for lizards. walie in Abyssinia. distinguished by the form of their horns but otherwise closely similar and still completely fertile with each other. raddei in the Caucasus. usually peculiar to the special districts. Tibet. of 52 amphibians and reptiles 16 were new. is effective. Caves. An unusual number of the species collected there by Whitehead in his expedition of 1886 were new to science. of the river basins of Borneo has its special orang. with identical environments. very characteristic fauna. such as the genus Carabomorphus on Kilimanjaro and the Gurui Mountain. 41 pairs of species are found on 78 the north and south sides of the Amazon. 111 Thus each Valleys are isolated by mountain ranges as mountains are by plains. C. and the beetles included also an unusual number of new and remarkable forms. Vicarious beetles are known from the mountains of Africa. with 4 new genera. ibex in the Alps. Thus. and Arinodromus on Kilimanjaro and the high 44 Mount Kina Balu in North Borneo has a plateau of Lisca in Shoa.

10.110 The Effect of Geographic Isolation ever. After Brauer. and these form subspecies confined to a single or to a few neighboring caves. and giraffes are split into a number of geographic races. The small snails of the genus Lartetia. high degree of differentia- A tion into species in cave-dwelling forms is the frequent consequence. The silphids of caves. are present in numerous places in the Jurassic and Muschelkalk region of southwest Germany. which are in part sharply defined and in part connected by transitional forms. and the species fall into geographic groups. 10). With great constancy of generic characters. 25 Of the* type of Lartctia corresponding to a special type of spring. but each of the four peninsulas has 7 its own local forms/' There is an endless series of geographic races of the Apollo butterfly Parnafssius apollo. each FK. in part also alive. The geographic variation in this butterfly is favored by an extraordinary individual . and acicula. subfamily Leptoderinae. from left to right: L. pcllucida putei. . which ranges from Syria to Finland and from Austria to beyond Lake Baikal. antelopes. Various species of iMiti'tia. up that almost every cave has its These represent a number of genera that are not very split Widespread species form local races and subspecies without special Thus the lions. are not so transitory as inland waters. they are very variable in minor features (Fig. The butterflies o^ Celebes are spread throughout the island.. 1 Beautiful examples of the effect of isolation in caves are presentee by the cave beetles in the limestone caves of middle and south Europe The carabid beetles Anophthalmus (a subgenus of Trechws) is represented by numerous species in caves. are so special species. sharply differentiated. in part as empty shells. zebras. the and snails is in caves distributed genus Zospeum pulmonate widely is broken up into 30 or 40 forms. isolation. They live in underground waters and reach the surface in springs. and they afford long-continued uniformity of habitat conditions. related to the Hydrobidae.

The specimens of a given locality re- semble each other like coins significant change stant difference. graphic races. rather than the degree of difference. Lack of wings and slowness of motion hinder its dispersal. the former devours the young of the latter. but even the most inmay be accompanied by a slight but con- have a similar behavior and distribution. The South American butterflies of the family Erycinidae are extremely restricted to their places of origin by a tendency to avoid flight. The erycinid genus Calydna differs from its relatives in the possession of strong powers of flight. where the two lizEuropean hare Lepus europaeus. and these are Planaria alpina is in competition with inhabited by L. Melinaea. furnish these conditions. and thus prevents its spread. genera. ten varieties of D.. elsewhere a mountain and arctic form. f til initiator are situation is The listed in central Europe. and in this genus the individuals of a species. alpina is found only in the uppermost parts of spring brooks. Other American Lepidoptera. L. of locality from the mint. Thus Lcpus twmhis. vivipara. some remnants have been able to maintain themselves in isolated areas when barriers prevented the entry of more advanced competitors. especially frequently exhibit no differences and are morphological distinguished primarily by means of coloration. . occurs at all levels in Ireland. the arctic territory. Of certain species. Constancy of a character in a series of specimens. and bogs and swamps. and Primitive Forms 111 similar with the goat chafer Dorcadion. At intermediate heights P. such as Heliconia. the geographic races are accordingly innumerable. especially in places without suitable sites for egg-laying or where the temperatures are too low to hatch them. of mammals and birds. even south of the Alps (for example. in Lombardy). so that numerous local races have arisen. vivipara occur together. ards Laccrta a^ilis and L. civipara is able to maintain itself only in areas where its enemy cannot live. which formerly had a wide distribution. Planaria gonoccphdla and Polycelis cormita wherever the water tem- perature has a range of variation of more than 6 C. even from widely 75 Such geoseparated localities. Mountains above 1200-1500 m. and Mechanitis.Isolation variability. and even orders. where it is free from the competition of the 71 In Germany. exhibit no appreciable variation. and primitive forms Another effect of geographic isolation is the preservation of groups from destruction by removing them from competition with their more advanced relatives. forms the basis for the recognition of subspecies in the practice of Isolation many American taxonomists. families.

. which has 79 Southern Australia and Tasmania harbor evidently replaced them. Australia affords one of the best examples of the preservation of archaic types. they represent remnants of stocks found nowhere else. to a few South American genera. survivors of the primitive crustaceans of the family Anaspididae. lives here. Schizopoda is likewise the home of certain primitive relicts. and to the North American opossum. Their primitive character is shown by their relations with the Palaeozoic genera Uronectes and Palaeocaris. number of languages is found on the slopes of the Andes. Paranaspides. primitive of all The most New Zealand and Australia are the home of the primitive mygalid spider Hexathele. The isolation of Tasmania has even preserved two species of predaceous marsupials. It is so closely allied to the caddisfly genus 70 Rhyacophila that it almost forms a link between the two orders. in some cases. and their intermediate position between the and Arthrostraca.112 at altitudes The where Effect of Geographic Isolation competitors are excluded by the uniform coldis its it ness of the water. Almost all the language groups of North America are represented in the relatively restricted area between the In South America an astonishing Pacific and the Rocky Mountains. which are extinct on the Australian mainland. Thylacinus and Sarcophilus. Groups speaking a large number of apparently unrelated languages are found in the Caucasus. together in Disappearing the human groups have been crowded much same way in unfavorable and inaccessible areas. Here are found the only egg-laying mammals. the sole survivor of a Sphenodon formerly widespread order of the most Liopelma represents reptiles. while the 84 rest of the continent has no more than a dozen language groups. with two genera. and in Recent deposits in company with those of the dingo. two large and widespread orders. It is rightly called the land of living fossils. primitive of the families of New Zealand is frogs. these are veritable "missing links." the surviving marsupials. Echidna and Ornithorhynchus. consisting of 3 monotypic genera: Anaspides. with their numerous reptilian characters. widely distributed. where their remains are found in the Pliocene. With them lives the greater proportion of which formerly ranged over Eurasia and North America but are now restricted to the Australian series. and Koo- nunga. languages that seem not only to be unrelated among themselves but also unrelated to other known languages. the hepialid Palaemicra calcophanes. There seems to be an accumulation of people representative of numerous linguistic stocks in such areas. Lepidoptera.

the physostomes. Pholeoteras. the great auk. of the cave snails of the Balkan Peninsula. When and foxes of the settlers. housecuts. rabbits. they promptly demonstrate their competitive superiority over primitive inhabitants. The three remain- The more ing genera of the extremely ancient lungfishes are fresh-water forms. as happens nowadays chiefly through the agency of man. Hydra and Microhydra and their relatives. the primitive types are When. The simple coelenterates. have thus been able to survive in it long after they became extinct in the sea. primitive bony fishes. including the proportion Underground caves are genera Meledella. long. introduced in the isolated areas. of a group whose representatives are otherwise absent in fresh waters in Europe. 8 Fresh water is an impassable barrier for many marine animals and Animals that were able to adapt themselves to fresh water plants. and Speloeoconcha. The endemic birds of Zealand give way before the buntings. the 2 "' 11 and many more are gone. In the same waters in which it occurs are found crustaceans of the genus Troglocaris. and later entered the such a protecting barrier is broken down. In all parts of the southern hemisphere the native earthworms disappear with the introduction of Lumbricidae. the dodo. the effects are still more The list of animals exterminated by thoughtless action is severe. is entirely unrelated to the present snail fauna of the surface and represents a remnant of a fauna no longer existent there. 88 Animals of subter- ranean waters are likely to be peculiar and primitive. make up a large fishes. The ganoids that dominated the seas in the Mesozoic survive primarily in fresh waters. Thus the marsupials of Australia are disappearing before the introduced cattle. majority of fresh-water Relatively few of the spiny-rayed Acanthoptcrygii have become adjusted to the fresh-water environment. Phygas. When man himself becomes an enemy. and goldfinches. the passenger pigeon.Isolation and Primitive Forms 113 A large also a refuge for primitive forms. The blind cave salamander Proteus anguinus has no relatives in Europe. and solitaire on Mauritius and Reunion. Bathynella natans. . modern animals are usually doomed. is the only relative of the above-mentioned Anaspididae found outside Australia. sheep. by accident or intent. found in deep wells at Prague and Basel. starlings. Steller's sea New cow Hydrodamalus. The array of archaic fishes in fresh waters supports the theory that fishes originated in flowing inland waters sen. and still others are in danger quagga. are probably preserved only because of their adaptation to the fresh-water environment.

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E. 149 ff. Vcr. "Studies on northern MichiEcology. ihre Ausbreitting. Ortmann. Evermann. pis. und Hausrind..116 37. 208. 4:1- 367. "Die Insel-Reptilien.. A. 33 figs. Jordan. "Bcschreibung von Ilybriden zwiZ. 40. 50. Mensch und Landschaft auf Celebes. 56." Zoologica. Mobius. Museum. "Kiifer 1-42. 1912. Breslau. 1-4. Minna D. Karl. Robert. Wc.. Rhltl. Lack. G. 1905. and Harold W. 1 fig. 1 pi. Brown. Leche. 33:72-75.. 65:151-162. Darwin's finches. Acad. Heft 2. und Netzfliigler. 2-4.. "The fishes of Samoa: Description of the species found in the archipelago. sclien figs. 5 51. Acad. U.sf. 16:1-48. illus. Fisheries. 1906. of the Rhio-Linga Archipelago: under uniform environment. 25:17-3-488. "Hybridization between Rana pipiens from Vermont and "Nachwort." 1916. Lendenfeld. 37:1-9. fishes of the 5. 2 text 55. 44. S. 56:1-79. Wisent. 47." Bull. 30). Cambridge. 54. 111 text figs. (pt. 47: pts. Iwanov. 10:427-475. Svenska Vetenskapsakad. Isolation Philiptsehenko." Siiugetiere": Wilhelm. pis." Bull Bur. Bull. 53." 1887.. Moore. 49.. Lonnberg.. Hirt: 95.. 1947. Abst. 1929. The Effect of Geographic and J.. 33-52. 52. "Mammals collected by the Swedish Zoological Expedition to British East Africa. figs. Kammerer. L." Kg/. U. Ann. 48. 42. "Check-list of Hawaiian land and fresh-water Mollusca. 2 B. Variation. "Die erdgeschichtliche Bedeutiing der lebemlen Verhandl naturforsch. 1907. 1947. H. 1915.." Evolution. 3 J. study of specific differentiation Natl. Matthew. siidpol Exped. Idem. Martin.). 1911. Handl. a S. 2:87-108. 23:504-506. D. 46.. J. David. Bishop Museum. Erne faunistische Studie. . pp. 10 maps. Natl. pis. "The Swedish South Polar Expedition. 1. S. 1898. 1-6. with a provisional check-list of the fishes of Oceania. 1898. S. (p. W. 36 figs. "The mouse deer pis. 9 figs. Wilhehn. und Chrysoehloridue." 7. Ernst. Kaum.. 1:263-288. A. 39." pi. v)." Proc. 1928. Paul. Robert von. 41. xiv. E. 4 pis. (p. "Die Siisswasser-Coelenteraten Zool. und Artbildung. pis. 1909. 1934. Cambridge Univ. 54)." Science. Tress: x. "Die Familien der Centedidae. Ifg." and W. Der Artenwandel auf Inseln-. No. 4 45. pis. eastern Mexico. 19:i-v (p. 4." Deittsch Oxtafrika. 8 pis.. pt." 24:171-318." Proc. E. 1935... Jewell. 6:1-69. E. Einar. 20:1-158." 57. D. Jordan. N. 4 pis. Vienna. "Phylogenie. and Alvin Seale. gan bog lakes. Australiens. "Fishes of North and Middle America-"" 149-153). Natl.-Lchrc. (pp. 1S96-1900. Jalirh. Sci. S?/. 1908. figs. Museum.. Bison. S. 4. Najadeen. Mcrtens. 84:1-209. Sci.oolot>ica (Stuttgart). "Climate and evolution. 4: No. 1906. 43. induct. 48. Y. "Zur Entwieklungsgeschichte des Zalm-systems der Teil 2. Kobelt. "A case of isolation without barriers. Deuticke: 324. 20 38. Mayr.. Kolbe. Deutsch Ost-Afrika. 5:1-188. schwed. 15 pis. Kornrumpf. Miller. 1947.." Ergeb. 1926. Solenodontidae.stf. "Ecological factors in speciation..

Leipzig. Calwers Kaferbuch: Einfiihrung in die Kenntnis 1907. Soc. David. 1 fig. 62:609-666. A. Camillo. 4. . 93:1-366." Fauna llawaiicnsis. edited with the assistance of well-known Stuttgart..Bibliography 58. Darwin and after Darwin: an exposition of the Darwinian theory and a discussion of post-Darwinian questions. Perkins. Nat. 611). 1898. Ribbe. 435). 27. including a monograph of the American species. 215. "Hyinenoptera aculeata.. Constable: xiv. G. 1897. Proc. Putnams: xv. 1932. 30: p. The shellfish of New Zealand." Ann." Cambridge Natural 626." Reise in Ostafrika von A. in Cow/io/. 65. Seit/. N.. 26 pis... Polinski.5-292 in Shureliff. 693). pi. 1900. Snethlage. G. 9 ff. 6:xii. 72. 1916." pp. 1912.. R. C. London. "Monographic der Daphniden Deutschlands und der A." "Non-marine Mollusca of Patagonia. F. "Ober die Verbreitung der Vogelarten Unterama- zonien. and Auguste Forel. Museum R. Pax. Pocock. on the zoogeography of the Pacific Islands. 1903:340-368." Deutsch Ostafrika. 1903. Monographic des Genus Bosmina." Bull V. Schaufuss. European animals: their geological history and geoLondon. 52:561-567.. der Kafer Europas. Carl. Sars.. 29: p. Humblot: viii. 60. J.. Bosmina benachbarten Gebiete: I. Idem. Jungle Islands. Zoo!. 2nd ed.. 20 pis. Petersburg. Pilsbry. 7 pis. 1912-1913. 73. (p. Smith. Natl Museum.. Schmidt. Adalbert. E. 117 1909. Christchurch.. Idem. 1 pi." 61 (p. 62. 1906 of all the specialists. 1913. A. Sci. Sharp. Zoologica (Stuttgart) 25. "On and Achatinella. Voeltzkow. p. 6. "The sessile barnaeles (Cirripedia) contained in the collections of the U. Syst. 7-8 (p.. (vol. "On some additional Crustacea from the Caspian Sea.. pis. 8:231-251. Lehmann: 14 figs. 63.. text figs. National Museum. Powell. 58- Wladislaw. Whitcombe & Tombs: 1-106.. Y. 1-2. 18:1-13. Scharfi." Zool Jahrb. . Patagonian Exped. don. 28 text figs. F. 6th ed. 67. 1930. O. St. 107 ff. "Die Aktinien der ostufrikanischen Inseln. 1:1-122. "On the geographical distribution of spiders of the family Mygalomorphae. vols. A list of the land 8. E. 1895-1899. 71. Ferdinand. W. Emilia. 1899. J. 1911. 84). 342). P. 1896. 76. 298. F. 1897." Ent. 78. 4th ed. 2:399-418. known Macrolepidoptera of the world: a systematic account Macrolepidoptera. Acad.. Ornitliol. 3:513-633. Princeton Unic. pis. 533). 74. 22-24.. 293 (vol. 64.." Rept. Scluuidinn. F. illus. N. vols. text figs. 5.. No. 1946." /. Sees. Hist. 01. (p." 141. "Die reliktare Gastropodenfauna des Ochridapis. 47-371. 61:469-539. "Anleitung zum Sammeln von Schmetterlingen in tropischen Landern. and freshwater mollusca of Trinidad. Peseliel.. R. H. S.. 69 figs. 2:27-3-305. coregoni im baltischen Seengebiete. Lon- 68. figs. I.. O. 258. 75. A. 5:81-584. 66. "Essay S. 2 pis. 1 map. 1913. Dunker & 1883. Phila. 59.. 77. pp. Rundschau. Sci. 76 pis. figs. the zoological position of Partula Proc. L.. 2nd ed. Romanes. K. Versucli einer Neue Probleme der vergleichenden Erdkunde als Morphologic der Erdoberflache. Ruhe.. S. p. Acad. 3 maps. Stuttgart. "Insecta. Longmans: 3 69. Heft 63:1- 70. "Rhizopoda. B. Zool. graphical distribution. Schweitzerbart: 2 \-ols. illus..

Congr. 113 pis. 1893. G.. "Geographical distribution and dominance in relation to evolution and phylogeny. Zacher. Wagner. Van Denburgh. Kin Bcitrag xur Geschichte dcr Siisswasserpis." Zool Centr.. Adolf. 3 81. 4 pis. 1917.: 82-821. 1933.S. Whitehead. Sinisa. Sitzber. Leonhard. Kl. Univ. 57 1 Galapagos Islands. fanna des Balkans. 93.sclilands und Fischer: 1-287 (p. 92.. Spencer. 91.-n. 1934. (4) 2:133-202.118 79. H. 206. 18:1-298. 1912. Sri. 1901. Introduction. F. U. J. 484). (p. 83. govina. Entomol. Natl Museum.. John. "On the fauna and zoological relationships of Tasmania.. 52 figs. E. 275). Occasional W. 88. Moriz. 317. 203-374. Jena. with notes on the iguanas of the genera Conolophus and Ainblyrhynchiis. 5 85. ihre Vcrhreitnn". Idem. "On the whea tears (Saxicola) occurring in North America. "Nouvelles contributions sur 1'origine du lac Baikal. "Uber die Verbreitung nnd okologir der (^urllrntricluden auf der Balkanhalhinscl. London. Anton.. [7. Adv." Proc. Gurney & Jackson: xi. Stcner. Stejneger. Nortli Borneo. "Hb'hlenschnecken a us Siiddalmatien und der Herce86. Schwabe: iv. 1948. 80. 294. 1913. Proc. sci. Abt. Entstehung der Arten durch ranmliche Sonderung.S. Vosmaer.). 1914. Calif. 1889. Wien (m. "The avifauna Papers Calif. Insects of Hawaii. 18: 694-696. 21 figs." ibid. 123. J. 1. Sri. 90. A. 1 of the figs. 305 (p. "The Porifera of the Siboga Expedition: II. 82. Basel." Zoogcographica. Wagner. 84. Verescagin. 1933. vol. "The giant land tortoises of the Galapagos Archipelago.. 23:473-481. 667. Exploration of Mt. 1:33-48. The genus Spirastrella. 87. 48-54 (in Russian). B.. John. Planktonkunde. 1910. 4th Meeting Australasian Assoc. "The Galapagos lizards of the genus Tropidurus.K." Rcpt. 32 pis. Leipzig. R... Die Geradfliigler Deut. So'. pi. . Wiss. 1892.. 1931. 723. C. S. Honolulu." C. Zimmerman.. 2nd Intern." Trans. G. Acad. 22). C. Stanknvic. 1914. Tenhner: \\. figs. Acad." 89. of Hawaii Press: xx. Taylor. Swarth. The Effect of Geographic Isolation W. S.. 1912." map.. 2:147-203. Ahad. Acad. Kina Bain. Gesammelte Aufsatze. Oxford: 271pis.

. The Chamberlin's further theories of the larger outlines of geological 119 his- . for example. plained by the changes in the earth's surface in the course of geologic Stratified rocks. which cover great areas of the existing continents. show that there have been extensive invasions of these land areas by the sea. on the evidence of the presence of submarine continuations and Norwegian river valleys to a depth of 60 m. require complete revision in the light of the much more adequate planetesimal hypothesis of Chamberlin and Moulton. 11 The invasion of tin* continents by relatively shallow inland seas need not affect the general permanence of outline of the great continental blocks. The fauna of southern Victoria agrees closely with that of Tasmania and differs markedly from that of northern Victoria and New South Wales. whose rhinoceros. can be shown to have been joined to them by dry It is generally accepted that Great Britain was land in the past. notably Great Britain and many East Indian islands. There are fre- Eurasia than to South America. based on the nebular hypothesis. where they are frequently brought up by fishermen. Conversely. numerous land masses now separated from the adjoining continents./ Historica I Zoogeography THE DISTIUBUTION OF LIFE IS present arrangement quent examples of areas formerly connected now entirely separated. North America. older concepts of the origin and evolution of life. geologically speaking. such as the of the British Dogger Bank in the North Sea. off Bremen. Many such facts are extime. has much more faunal resemblance to of barriers NOT SATISFACTORILY EXPLAINABLE BY THE and land connections. on the limits are defined mammoth and by the continental shelves rather than by existing shore lines. united with the European continent until recent times. and presence of the presence of the remains of large land mammals. of the of terrestrial deposits at a depth of 200 m.

Permanence of barriers Mountains have been no more permanent than marine barriers. The periods of uplift are marked by formations. and cycles. The deserts of the Great Basin in North America are in part the bottoms of extinct lakes. may often be shown to be the remains of once higher ranges. that suggest arid conditions. even than some species. were dcmonstrably uplifted in the Tertiary. 51 Present-day records indicate a climatic change to warmer and drier summers. The very ranges that are now highest and most important as distributional barriers. the earth has passed through a number of climatic the extreme of humidity. It is legitimate to suppose that the this region . whose importance as barriers has been reduced by long-continued erosion. these periods culminate in great extensions of glaciers from mountains and from the poles. such as the American Cordillera. Steppes and deserts also change. which corresponds to base-level erosion of the continents. The uni- form base leveling corresponds to widespread deposits of limestones. Isles and the Faeroes. Some mountain ranges are younger than many types. ending in coal formations. accompanied by extensive transgressions of the continental borders by shallow inland seas. perhaps partially induced by drainage and stream regulation. and both the land and marine fauna of this area would be powerfully affected.120 Historical Zoogeography tory are of special importance to historical zoogeography. they are formed anew or are recaptured for a renewed development of plant and animal life. the Himalayas. From ment zonal climates. There this was are traces of a recent steppe period in central Europe. and the whole series of successive ranges from central Asia to the Alps and Pyrenees. the cycle passes to the opposite extreme of cold. The impermanence of fresh-water basins and of rivers has already been discussed. area/' with disappearance of forests and a return to the steppe in this The ruined cities of the deserts in central Asia indicate that was inhabited at a relatively recent date. According to these theories. on the other hand. often red in color. and uniformity. with accompanying uplift of the continents in adjustto the isostatic balance and with renewal of erosion. aridity. the Gulf Stream would be deflected from the Scandinavian coast. brought to an end by conditions favoring extension of the forests. Climatic changes have gone hand in hand with the changes in the Should a submarine ridge arise between the British earth's surface. Low mountain regions. of animals. warmth.

to be found in the in the south polar region is indicated by On the the discovery of silicified tree trunks in Kerguelen Island. we must assume that. was covered by a continuous continental glacier. may depend on or barriers. The presence of coral reefs of earlier geological epochs in northern latitudes also indicates a warmer climate of the condriven out by a temporary seas. less protection from sunlight. water temperature of 20 C. it is evident that the past rear- rangements of land and sea and of other connections and barriers must have been of profound influence in the dispersal of animal life on the earth s surface. while their with lower temperatures. are found in the north polar regions. The most recent period of glaciation was divided into interglacial periods during which animals intolerant of cold invaded northern regions. precipitous topography. spaee/'- Satisfactory explanations of present faunal relations the recognition of the presence of former connections as well as with Zoogeography must reckon with time On the basis of evolutionary theory. even in the tropics. of the present. during which a large part of the northern hemisphere was covered by glacial ice. In Europe the glaciers of the Scandinavian mountains and of the Alps extended far into the plains. These remains indi1S cate a warmer and more humid polar climate in the early Tertiary. Traces of glaciations in much more remote epochs may be observed in many regions. only to be new advance of the ice sheet. and more violent winds. Changes in climate have also taken place independently of topographic changes/ Fossil remains of dense forests. for reef corals are now restricted to the tropics require a minimum Iceland and the Arctic of relatively recent periods exhibit a fauna found only far to the south. other hand. The climate of the lowlands was wholly different from that A large part of North America. and glaciers were present even in the intermediate ranges. such as are now warmer parts of the temperate zone. there is abundant and conclusive evidence of the existence and similar climatic change of a glacial period at the close of the Tertiary. where the length of the winters and the dryness of the air now make tree growth impossible. and Marine deposits of now have a direct Since barriers are important for the evolution of animal forms and effect upon distribution. accidental . New mountain ranges influence the amount of upper levels. offer wholly new conditions.Permanence of Barriers and Pacific 121 Tertiary opening of the Central American straits between the Atlantic rents Oceans must have had far-reaching effects on ocean curand climates. at the same time. precipitation in adjacent areas.

122 dispersal Historical Zoogeography or extreme vagility aside. is Parallel evolution is indicated in some supposed also maintained. genus. in Nevertheless. for the various connections that have influenced the dis- Forms persal of the group need not have been contemporaneous. derived from the marine Cerithiwn. It must have arisen 1 The* same here from ancestors related to the Polynesian Partnla. and neither living nor fossil The forms of this group are to be found elsewhere. of holds Hawaiian for the endemic situation birds. The is snail California) genus Potamides of brackish water (India. whether speeies. Thus the numerous species of snails of the family Achatinellidae are confined to the Hawaiian Islands. the family general Drepanididae (cf. Such continuity is primarily a continuity in time. must inhabit a continuous area. borcalis appears in the 1 genera. that the two populations of hor calls borealis have been independently derived from the tropical form. or family. The sea is not a barrier for them. In the sense that different ancestral forms could not give rise by convergence to identical descendants. for the number of separate char1 would have to fall in line arise is unthinkable. this acters that is unquestionably valid. with a common ancestry must of necessity have had a common center of origin. Another example of a natural group with continuous range is afforded by the penguins. Africa. though 1 . identical varieties localities if may widely separated they arise as modifications dependent on the influence of The widely distributed appendiculate l : from the same stock the same conditions. It is much more probable that different Cerithiwn species have developed into the various Potamides species under the influence of brackish water in the different parts of the world than that all the species of Pol (nn ides are to be derived from a single ancestral form. There are certain modifying corollaries to be added to this basicprinciple. center of origin is sufficiently obvious in many groups of animals. every natural group. The contrary hypothesis. It would perhaps be better not to speak of the genus Potamides but to refer to Polamidcx forms of the various Cerithiwn species. this statement applies equally to groups of higher rank. Wallace states that "two identical species have never been developed independently in widely separated areas". Chapters 6 and 26). rittilar'ui same subspecies (recognized as the forma tijpica) in both the north and south polar seas and may be the persistent ancestral type from which the form in intermediate seas is derived. and. or an area that has at least been continuous or connected at some time during the history of the group.

whether in consequence of " 11 1 alteu d"TiaI5itaF^conditions or specifically through competition with more advanced forms. The range of the Pyrenean shrew Myogale pyrenaica is widely separated from that of its relatives in the steppes of southern Russia. The ranges of families and genera and even of species are often broken by broad areas in which the forms in question are absent. The varying hare Lepus timidus is represented in Ireland and the Alps as well as in north Europe and Asia. usually of the same species. " r> Such continuity of distribution is by no means invariable. and Germany. the Pyrenees. horse. and jerboa. These instances. but its northward spread has been favored at this point by the cold waters of the Humboldt Current. DjsmnHnnifry of range is accordingly explain! by extinction in the intermediate areas. is the rule for the higher groups. gopher. Subfossil bones testify to the former presence of the ptarmigan and alpine hare in central Europe. A whole series of arctic animals (and plants as well) are to be found on Mount Washington in the White Mountains of New Hampshire. in which there are isolated outliers of an otherwise arctic range. Their fossil remains are likewise known only from the southern hemisphere.Permanence of Barriers 123 their headquarters are on antarctic and subantarctic shores. where it has presumably become extinct in consequence of changes in climate and vegetation. One species has New reached the Galapagos Islands on the equator. One may even say that discontinuity for genera. even perhaps but certainly for families and orders. the greater will of probability discontinuity of range in the modern survivors. are amply accounted for by the southward extension of a cold climate in the glacial period. in alluvial defollowed the glacial period posits make it probable that a steppe period . and climatic changes may The the from inferred in composition and distribution of faunae. The more remote the* origin of the related forms. live in Labrador and Greenland. but fossil remains of this genus are found in the terrestrial deposits of France. Their relatives. England. they withdrew to the north and to the higher altitudes of the mountains. wide distribution abundance and steppe The "eon verse reasoning such as the saiga antelope. with the retreat of the glaciers. a few species range north to the southern tips of the three southern continents and to the South Island of Zealand. Belgium. of the remains of animals. i is also possible. are numerous instances in which a former continuity of range period of common be the There can be demonstrated to explain the present discontinuity. and the Caucasus. the ptarmigan Lagopus mutus has representatives in the Alps.

As has been stated previously. was a member of a flourishing family in the Mesozoic but is now the sole survivor of the tetrabranchiate cephalopods and occurs in only a few species in the Pacific and Indian oceans. He explains this area. The changed conditions may not be and every kind of habitat factor may be involved. i. such as that of the shrew. There are relicts from periods when the competition in an animal community was different. although the evidence for the existence of such a steppe period is far from being as abundant or conclusive as that for Such distribution of steppe conditions in the past would explain numerous instances of discontinuous ranges. where they have escaped the brunt of the struggle for existence with the more modern bony fishes. marine relicts. better adapted climatic ones. appear as anachronisms. In a limited arid area in Moravia. and one speaks of glacial relicts. for example. This is commonly true of relict forms in the sea. The rise of new competitors. in the changed environment. than the older forms to engage in the struggle for existence. 31 covered only with a little coarse grass and a few stunted bushes. glaciation. and the few that sur- whether by isolation or by exceptional ability to meet competition. The genus Nautilus. which have supplanted the ancestral Mesozoic ganoids in the sea. a relict of the steppe period/ evidence from reptiles with Great Plains affinities in the central United 1 *1 States indicates a similar postglacial eastward (instead of westward) extension of a grassland fauna in North America.. The few modern ganoid fishes have become adapted to fresh water. Myogale. Burr found more than eighty species of Orthoptera that were wholly unlike those of the surrounding territory and exhibited a high degree of resemblance to those of the Volga valley. steppe relicts. Whatever the reasons for the vive. as in the glacial and steppe relicts.124 in central Historical Zoogeography Europe. 10 satisfactory explanation of the discontinuity of distribution of certain animals by demonstrable changes in the environment. as a faunal island. prob- The ably correctly. occurrence of relict groups. makes it possible to conclude that in other instances present discontinuities are based on changes in a former The . 1 '5 when habitat conditions and faunal relations were different from those of the present. relicts. are termed relicts.e. leads to the destruction or dispersal of the older forms. The few modern genera of pentacrinids are confined for the most part to deep seas and represent a group that flourished in the Jurassic and Cretaceous. they indicate that some sort of change in their physical or biotic environment has taken place. mentioned above. such survivors of a former period. etc. where the physical conditions have undergone relatively little change.

The number of genera in common is large.{ ' :ir> . or extensions of the ocean. some of which are now extinct in one or both areas. in spite of the fact that the parts of the sea that separate them are impassable barriers for the larger animals. moose.-*' This land bridge probably existed at Bering Strait and across Bering Sea and may have included part of the Arctic Ocean. and the species of the same genus are frequently so closely allied that they may be regarded as subspecies of the same form. and bears are thus closely related. the littoral fauna of the and Atlantic coasts in this region exhibits a large number of genera of fishes and invertebrates in common. same role Another explanation of bicentric distribution is extinction of forms from the intermediate area. climatic changes. Though the isthmus of Panama now forms an impassable barrier. and within these the peculiar phenomenon of "paired species. along with the Eskimo curlew. Although these populations seem to have no opportunity to meet and mix. The shallow seas in this region favor this hypothesis. In respect to marine life. a Missisout. such as mountain ranges. rise of superior competing forms. Pacific . bodies of land is separating formerly continuous parts of the sea play the in producing discontinuous or vicariating distributions. 125 One explanation of such discontinuity of related groups that barriers to dispersal of some sort have arisen.Permanence of Barriers continuum. the lynxes. and this hypothesis is supported by geological evidence. while the wisent and bison. found at times in the There are at present two distinct populations of the Hudsonian curlew breeding. sippi valley It is group. and wintering on opposite sides of the Americas. An example of similar faunal relations is presented by the marine faunae on the two coasts of Central America. reindeer. This resemblance extends to such Tertiary animals as horses and camels. in the period since the early nineteenth century." the geminate species of the direct relative of a given species being one of the Jordan. The great similarity between the and of that central Europe leads to the confauna of Great Britain clusion that a land bridge existed here also. migrating. and the various deer are only a little more distinct. deserts. The beaver. 57 The mammalian faunae of Eurasia and North America exhibit a high degree of similarity. was wiped possible that the connecting link. and their separation seems to have been comparatively recent. The supposition that a land connection existed between the two continents in geologically recent times and that such a connection must have existed at various times in the Tertiary thus acquires a high degree of probability. no racial distinction has been detected.

which increases the reliabilThe groups examined are distributed as ity of the conclusion.9 . probably geological evidence supports the zoogeographic indications. Similarity (i. may be supposed analyzed the fauna of Celebes. example.e.(i phibians Birds 37.!)/) "28.8% amI7. the connection continuing at most into the early Eocene. The two continents are thought to have been connected in the Upper Cretaceous and perhaps in the Paleocene. The subject is reviewed by Ekman. which are taxonomically also have decidedly different ecological relations. especially the mollusks.(> .(>' (>.who joins phenomenon There is are. Ki. for 1 in the opinion that all this evidence indicates former broad water connections between the Atlantic and Pacific in the Panama region and perhaps This is also elsewhere across Central America.* 11. their evidence it appears very probable that the present fauna has Sarasins island in the Eocene.9." M The Eocene to in full differences Mid-Pliocene. and birds. agreement with the explanation of the deep-seated between the Tertiary mammalian faunae of North and South America as resulting from long isolation of the South American continent. amphibians and reptiles.>. follows: Widespread Forms and Uncertain Kn- demic Species Mollusks Reptiles and 30. Jordan demonstrated at least 100 such pairs of species of fishes.1 !<). Separation extended from to late Mid -Pliocene. These examples show how both periods of union and the appearance of separating barriers may be dated by the comparison of the early 1 ' geologically older elements of the faunae of given areas.> <. The 42 reached Celebes by four distinct routes since the emergence of this The groups investigated. and distinct widely exhibit surprisingly comparable relations.0 Average 28. with reference to relations On the basis of with the inhabitants of the surrounding islands. differences to have arisen since the date of separation.O 10.>. .126 Historical Zoogeography opposite coast instead of some congener on its own coast. 1">.3 8.3 *!. 33 genera of decapod crabs listed by Rathbun as confined to tropical American waters and represented in both the Pacific and Atlantic..8 . and the clearly represented in various groups of invertebrates. homology) of faunae speaks for union at the period of origin or active dispersal of the forms in question.

Permanence It is of Barriers 127 and of the mammalian fauna the remainder. inasmuch as geologic evidence in this F. are wanting. The value of zoogeography for the elucidation of former land connections seems to be the higher. and. which reached Pennsylvania. deduced in arras now continental. Conclusions as to former land connections and former climatic arrangements there is thus be based upon zoogeographic data. now separated by seas. to palaeogeography. specifically may be established and dated with some degree of Zoogeography thus becomes an important aid to geology. evidence. if abundant palaeontological evidence. from the correlation of are less convincing. so that only rarely. 40 Most refers to zoogeography as a form of submarine geology. in particular. which have left their deposits Land connections. An exchange of forms took place at this time. The older Tertiary rocks of South America contain remains of marsupials. Sarasin even direction is unavailable at the bottom of the sea. arations. the from identity or differentiaPalaeontology. is of much later origin than Tertiary separation of this continent extended through a period of most active evolution of the mammals of the northern hemisphere. 1 hypothetical former land connections are based on the present distribution of animals/'-* The probability and possibility of the conclusions derived from these premises must be examined with especial care. and The platyrrhine monkeys. and llamas. similarly evident that a part of the fauna of South America. Palaeogeographic maps should be based on geological and palaeontological. It is evident that the latter series emigrated later and arrived after the rc- establishment of the connection with North America. is this kind of evidence at all adequate. tion of fossil faunae. for the armadillos and porcupines of North America are likewise relatively late arrivals. . peccaries. is most certain. but these three elements are of very unequal value. but remains of the present South American carnivores. contemporary with an invasion of ground sloths and glyptodonts. and of tapirs. as well as nco/oo' logical and neobotanical. as in the question of the former connection of stratified rocks in areas North America and Eurasia. and many have regarded this form of distributional study as the capstone of the zoogeographic structure. The geologic evidence of the existence and extent of former seas. such changes and re- may arrangements certainty. edentates. deer. is sometimes able to conclude with a high degree of probability on the nature of the contemporary connections or sepThe data of palaeontology are meager for the most part. hystricomorph rodents.

geologists are not at all agreed on the fundamental question of the permanence or impermanence of the continental land masses. phenomena When employed alone. the to geologically recent have a high value. Such critical care has only too often been wanting. they are of doubtful importance.128 Historical Zoogeography The principal and often the only source of evidence for the former existence of land connections remains in the data of zoogeography. Dana was the and oceanic areas were essentially permanent. and land connections were invoked to explain similarity of geologic structure or faunal resemblance even {l when they involved broad oceanic areas. after weighing the evidence on both Wallace " assumed a fixed sides. From Forbes through von Ihering and Scharff to Uvarov. Asaccessory evidence. leaves the question undecided. and even a ls large amount of evidence must be interpreted carefully and critically. 10 In spite of the repeated warning of conservative investigators. The unbridled of animal distribution <VJ hypotheses concerning the rise. The? controversy over this question is by no means at an end. and connection of land masses have left scarcely a spot which has not at some time been involved in a land bridge. In view of the numerous changes in the shore line of present land areas in former times and of the widespread presence of marine deposits on the continents. Dacque. R. and this field of zoogeography has become a clearing house for fantastic combinations. however. Suess 7 the opposing camp.7 defend the permanence of the continents (with the inclusion of the r and Haug 17 represent continental shelf) and of the oceans. A. displacement. The first to propose the theory that the continental ' r> . and this is plainly an inadequate view. with the statement that the distribution of plants and animals could he explained without supposing radical changes in the extent of land and sea. oceanic depth of 100 fathoms as the maximum over which former land connections were possible. there was at first no objection to the assumption that any desired part of the ocean might have been occupied by land. but this evidence is least reliable and becomes progressively less useful as the period of the supposed connection becomes remote. some zoogeographers continue to 11 tinents as easily as a cook makes pancakes" (Darwin). Wallace supported this position from the zoogeographic side. Jeffreys -~ and Matthew . "make con- The opinion of geologists should carry the greatest weight in the reconstruction of former land connections. especially with regard changes. to support and illustrate the conclusions of geology and palaeontology. Unfortunately. the of animal and to facts distribution assumexplain plant tendency by ing the presence of land bridges has been uncontrolled.

tions within the The presence Australia. in the reconstruction of former land connections.e. the 13 33 Madagascan Mantclla from an African ranid ancestor. beetles. emus and cassowaries in and the recently extinct moas in New Zealand. center of dispersal accordingly has reduced value. and they This evidence for an antarctic that these forms belong to a natural are now placed in distinct orders. together with the faunal relations. ocean journey in driftwood. Every phylogenetic by abundant palaeontological evidence. and the width of the intervening ocean must be taken into consideration. and these become less dependable the greater the degree of difference between the forms that are compared. was regarded as important evidence of the connection of these regions by an antarc-" and others. they inhabit Madagascar and tropical America. tic continent by Ilutton group is highly dubious.Permanence of Barriers 129 whole geological structure of the area. whose components are "homologous. has shown that there are numerous differMore exact investigation ences between the two groups. The must be interpreted with discretion in zoogeogbest evidence for former land connections is afforded by . the direction of submarine ridges. however. is dependent on assumptions. and they are now regarded as the and their wide separation in "relicts" of a 2 formerly widely distributed primitive type of insectivore. zoogeographic imon by active flight like birds." i. 5 It was formerly believed that the shrew-like Centetidae of Madagascar were closely related to the Solenodon of Cuba and Hispaniola.. The assumption. space was accordingly highly remarkable. such as snails and the pupae of longicorns and snout raphy. and the views of specialists on the rela- conclusion. The first step in the comparison of related faunae. or passively like spiders and small invertebrates with resting stages. taxonomically comparable. ostriches in Africa. is to determine the degree of relationship between the animals in question. are relatively unimportant in Relations between animals that are likely to survive an this respect. whether land connections. frogs formerly grouped together as the family Dendrobatidae were separated from the true frogs on account of their lack of teeth. The absence of teeth is now regarded as convergence. This is not always an easy problem. of large flightless birds in the southern hemisphere. bats. The - Different groups of animals are unequal in portance. especially with reference to the evidence they provide Animals dispersed through the air. and insects. unless supported same group may be divergent. the neotropical Dendrobales is supposed to be derived from a neotropical genus such as Prostherapis. rheas in South America.

130

Historical

Zoogeography

groups such as the Amphibia and most earthworms, to which salt water is fatal, and by non-flying mammals, because the transportation, even by large rafts, of such large animals as ungulates is impossible;
to starve
rafts.

even small forms such as mice, shrews, and squirrels would be likely on any rafting journey unless they were transported on sizeable

Mammals and birds have the great advantage for zoogeographic studies of relatively rapid and relatively recent evolution. Although the crustacean genus Apus, which is still living in fresh waters, was
already in existence in the Triassic, and most living genera of freshwater mollusks and insects are represented in the Eocene, and even the reptiles have numerous Eocene genera that still persist,* none of
the living genera of mammals are present in the indeed in the Miocene. The Pliocene genera of
the

Eocene and very few

same

though

mammals are largely as the Recent, but nearly all the species are different, 80 to 95* of the species of Pliocene snails are still extant. The

reasons for this

more rapid evolution of the warm-blooded animals are doubtless to be found in the general acceleration of their life processes, including the appearance of mutations, and in the greater
probability of the survival of new forms on account of their adaptability to environments that are closed to other groups of animals.

between related groups of mammals thus gives a clue to the amount of time since their origin, and consequently, in some cases, to the date of union of land areas now separated. Thus the presence of so many genera of mammals in Eurasia and North America represented by closely allied species or even subspecies in the two regions is the basis for the belief that these regions were

The degree

of differentiation

Even a separation since connected in geologically recent times. the Pliocene, from the evidence of the extinct Pliocene forms, would have resulted in a greater differentiation than exists.

The rapid transformation undergone by the species of warm-blooded animals explains the fact that the faunae of Madagascar, South America,

their insects or

and Australia are so much more peculiar in these groups than in amphibians and reptiles. Celebes, as an example, has no endemic genera of land mollusks, 3 among the fresh-water mollusks, 1 for amphibians and reptiles, but it harbors 16 peculiar genera of birds and 6 of mammals. Finally, the fossil remains of extinct forms are especially abundant among the mammals, more abundant at any rate than among other
*

The

lizards

Chamaeleo, Agama, Iguana, and Lacerta, and the

turtles

Emys,

Testudo, and Chelonia.

Permanence of Barriers
terrestrial

131

animals.

reliable information

Study of this palaeontological evidence affords about former connections of land areas, and some-

times also of separations, as was illustrated above in the discussion of the relation between the North and South American faunae.

Other animals besides mammals furnish similar evidence of former changes in the distribution of land. Land connections between areas now separated but inhabited by related animals need not have been direct; they may involve other areas and may have been successive rather than continuous at any one time. The more ancient the groups

whose discontinuous distribution requires explanation, the greater are the possibilities for their emigration by roundabout routes. The occurrence of two genera of mantids, Liturgousa and Stegmatopof animals

Madagascar and South America and nowhere else, and the example in the order Insectivora of the Centetidae and Solenodontidae in Madagascar and the West Indies, cannot be supposed to indicate direct connection between South America and Madagascar. Allied fossils have been found in the North American Eocene.
tera?
in

4

similar

Numerous examples

are

known

of

modern discontinuous

distribu-

tions that are connected

by a wider

distribution of fossil remains.

The camel

family

is

now

represented in Asia by true camels, and in

South America by the llamas and their relatives. This group had representatives in North America from the Eocene to the Pleistocene,

and

this affords a satisfactory

The pleurodiran

turtles are

now

explanation of the present discontinuity. confined to the southern hemisphere,

where they are widely distributed, but fossils of this type are known in Europe from the Upper Triassic to the Miocene, in Egypt from the Eocene to Mid-Pliocene, in India in the Lower Eocene, in New Zealand and in the Americas in the Upper Cretaceous, and in the South American Eocene/'"' The North American Cretaceous representatives of the genus Podocnemis were clearly inhabitants of marine 47 Direct connections between the southern continents coastal waters.
1

are accordingly not required to explain the present distribution of the

palaeontological evidence is have been adequately studied, a similar explanation seems to apply to the widely distributed families and genera of fishes, amphibians, and reptiles, whose present distribution is often a mere relict of a former world-wide range. Among reptiles, for example, the discovery in 1943 of the skull of an amphisl5 throws new baenid lizard in Lower Oligocene deposits of Mongolia light on the disjunct distribution of these limbless forms whose prin-

side-necked
available

turtles.

Whenever good
relations

and when the

cipal representation

is

now

in Africa

and South America.

132

Historical

Zoogeography

Although fossil remains of insects and spiders are relatively scarce, these groups also afford instructive examples of a more extensive distribution in the past, with a discontinuous or restricted distribution at
the present time.
Africa,

The genus

of ants, Oecophylla,

now

has a species in

one in India, one in the East Indies, and a fourth in Australia and in the Aru Islands; but no less than four species are known from the European Tertiaries, and the genus was doubtless widely distributed in Eurasia. Another ant genus, Ectatomma, with numerous species in the tropics of America, Asia, and Australia, is represented by a species in the Oligocene Baltic amber and by another in the Miocene Sicilian amber. A third genus, Macromischa, now has two species in West Africa and many in Cuba and Mexico; five species are known from the European amber. 16 The related lucanid genera, Lamprina and Neolamprina in South America and Sphenognathns in Australia, are connected by an intermediate form, Palaeognathus suecini, in

the Baltic amber.

Termites, represented in the tropics and subtropics by about 2000 now scarce in the temperate zones; two species reach southern Europe, although several live in the same latitude in North
species, are
Fifty-six species of termites are known from the North The primitive spiders with a American and European Tertiaries. three with survive abdomen genera; the group is represegmented sented in the European and American Carboniferous. 43 There are numerous other examples of this tendency, and their number is likely

America.

be greatly increased with the progress of palaeontological research. Although the incompleteness of the palaeontological record prevents the general application of this method to zoogeographic problems, the fact that gaps in distribution are frequently reduced by fossil forms
to

indicates that a similar explanation of discontinuity of range

is

often

to be invoked, even if the palaeontological evidence is negative. With the exception of small volcanic coral islands, such as those of Micronesia and Bermuda, and volcanic islands, such as St. Paul and those of the Hawaiian and Polynesian archipelagoes, there are perhaps few land areas that have been continuously isolated from all others.

This statement

is

undisputed for islands near the coasts

like those of

the Mediterranean, Ceylon, Formosa, Tasmania, New Guinea, and even Japan. The Indo-Australian Archipelago and the Melanesian

from New Guinea to New Zealand were probably mutually connected and thus also joined with Australia and southislands extending
*

A. E. Emerson, personal communication.

Permanence of Barriers
east Asia,

133

though these connections were by no means contemporaneous or of equal duration. Zimmerman 6 " reviewed this situation as follows: as a result of geological study, it is known that the Pacific Basin is underlaid basaltic rock; that in the continents generally lighter or

by heavy metamor-

phosed rocks

rest on heavier underlayers; and that continental shelves extend for various distances under shallow coastal waters and then

terminate rather abruptly at the edge of the deep water, which is of remarkably uniform depth of about 12,000 to 15,000 feet and more in the true Pacific Basin; that the volcanoes now above sea level have

from great depths and have been built of basic rock, and that no islands in the central Pacific Basin have true continental rocks been found. So far as is known to geologists, the only pre-existing exrisen
in

florae

tensive land masses in the tropical Pacific since the rise of modern and faunae are those west and north of Australia and on the

axis, possibly extending eastward to near the Tongan Trough in the neighborhood of Fiji. There is no geological evidence to indicate the existence of any large

New Guinea-New

Caledonia-New Zealand

land masses east of Tonga and Fiji. Africa was connected with Europe at times via the Strait of Gibraltar and probably via Sicily and Italy as well. Repeated connections

between Eurasia and North America must certainly have existed, probably across the present Bering Strait and parts of the adjacent seas. This bridge must have been in existence during a warm period, when the Alaskan climate permitted the dispersal of numerous animals. These connections seem to provide for distribution to all the regions inhabited by animals whose distribution is rigidly dependent upon the existence of land. The West Indies, also, may have been anciently united with each other and with the mainland, and Mada27 gascar was perhaps attached to Africa, though Matthew in 1915 the West Indies are defended the opposite view, that Madagascar and
strictly

oceanic islands.
is

more and more reliance on an explanation of the arrival of the foundation stock of immigrants even on isolated oceanic islands. The bulk of the insect faunae of mid-Pacific islands appears to have developed from wind-borne individuals. Over-water transport by flying birds is also a contributing factor, and floating natural rafts or even
There
a growing tendency to place
aerial transport in furnishing

Zimmerman,

driftwood provides another possible means of accidental dispersal. after much collecting on diverse scattered archipelagoes of the Pacific, believes that, although evidence for the former exist-

134
ence of a land mass

Historical
is

Zoogeography

lacking for that region, it is probable that high in islands existed there past ages in addition to those now present. Such islands could well provide steppingstones for the dispersal of
biotic elements out

even to the well-isolated Hawaiian Islands.

For-

merly the nearest high island was probably only some 500 miles from Hawaii in contrast to the 2000 miles by which they are separated
today. This hypothesis breaks the need for postulating thousands of miles of above-water transport, if land bridges are rejected, and intro-

duces the more probable hypothesis of a series of shorter over-water
66

steps.

the generally accepted connections are drawn upon a map regardless of their non-contemporaneity, the result is a connected
If all

land mass in the northern hemisphere with three great southward
projections.

between the southern continents through the antarctic land mass must be regarded with suspicion, although such connections have repeatedly been proposed and defended with ability. Many authorities, however, have found no inTheories

about

connections

superable difficulty in the way of a derivation of the southern faunae from the general northern land mass, and as long as this is possible and in the absence of direct evidence to the contrary, the assumption
of direct connections

be avoided.

27

There

certainly to are theoretic considerations that favor a derivais

between the southern continents

tion of the southern faunae from the north, based

ena of climatic change.

Land

on general phenombridges that extend right across thi*

Atlantic or Pacific are so dubious even from biological viewpoints that they deserve no consideration. The ambitious attempt of Arldt l does not carry conviction to a critical reader on account of its failure
to

weigh the evidence

for the

permanence of continental outlines

against the probability of transoceanic land bridges. Wegeper's hypothesis of continental drift is not in accord with
geological observations and not only is not needed to explain zoogeographical distribution but actually creates more difficulties than

many

it

61

adjusts.
lla

marized here.

The extensive literature on this subject cannot be sumand The interested reader should consult Wegener
0<i * tt

Du

Toit for the pro-drift argument. So far as the problems of animal geography are concerned, the weight of American opinion is
53 strongly against any hypothesis of continental drift. The distributional relations of the terrestrial animals strongly support the assumption that centers in the northern hemisphere have been

the places of origin of the principal advances in organization in the

Primitive Southern

Forms

135

more important groups of terrestrial forms. The vast land areas of the North Temperate Zone afforded a spacial basis for their development, and the successive periods of cooling and other climatic changes produced a periodic severity of selection and thus favored advances The groups whose adaptations or changes reprein organization. sented advances were enabled to enlarge their range and extend it to the south, at times driving more primitive forms before them. Successive impulses, following one upon the other, would push the earlier forms farther and farther to the south. Here and there, under
the protection of specialized habits or on account of barriers, remains of the primitive forms would be left behind as "hanging relicts," but in the main they would be driven to the three southern land

masses via present or geologically attested land connections. They would be preserved most effectively in these cul-de-sacs when the

appearance of barriers prevented the later, more advanced, comThis isolation is especially notable in petitors from following them. It is not at all Australia, which has been cut off since the Jurassic.
implied that advances in organization took place only in the north; favorable modifications may have arisen and adaptive radiation has occurred in various other areas, but the conditions for advance were

most favorable
cerned.

in the north, as far as terrestrial

animals were con-

Primitive southern forms

respect,

There is a great accumulation of evidently primitive and ancient " Australia leads in this vertebrate types in the southern hemisphere/ with its monotremes and marsupials and with the small
1

rodents whose only relatives exist as relicts on the mountains of 63 South America has its primiCelebes, Borneo, and the Philippines.
tive marsupials, edentates, hystricomorph rodents; its tinamous among birds, and iguanid lizards. Africa south of Sahara has preserved primitive mammals such as the tragulid dwarf deer Hyaemoschus, the lemurs, the aardwolf Proteles, and the golden mole Chrysochloris. In Madagascar the lemurs and centetids are primitive forms; the civet
is intermediate between other genera and, as such, is bird genus Mesites seems to be the most primitive memprimitive; the 65 Primitive groups of invertebrates ber of the crane and rail group. southern are also especially abundant in the hemisphere. Among in-

genus Eupleres

the most generalized type of termites occurs in Australia; the simplest Lepidoptera, resembling Trichoptera, are found in Australia and New Zealand; and about half the Australian bees belong to the
sects,

136

Historical

Zoogeography

58 has shown that the most primitive primitive genus Prosopis. Taylor in the southern continents and in land snails, Helicidae, are found New Zealand, and that progressively more advanced species are

found in the north. The whole mollusk fauna of South Africa may be characterized as a primitive one. The distribution of earthworms
is

similar in

its

arrangement, with primitive forms in the southern
in the northern.

hemisphere and advanced forms

Discontinuous distributions, with the most varied arrangement of the respective isolated areas, are the rule in the southern land masses.

The most primitive of the living odd-toed ungulates, the tapirs, are found in Malaya and in Central and South America. The dwarf deer (Tragulidae) are found in the East Indies, in southeast Asia, and in West Africa. The relatives of both of these groups are well represented among the North American and European Tertiary fossils. The iguanid lizards have their headquarters in South America and southwestern North America, with a single genus in the Fiji Islands and two in Madagascar. Their fossil remains are found in the Euro67 The giant water bugs of the pean and North American Eocene. genus Belostoma are widely distributed in America, Africa, southern Asia, and Australia. An isolated species in Dalmatia testifies to the former continuity of this range, and fossil belostomas are known from the European Miocene and Jurassic. 16 Hanging relicts, like the group of spiders with segmented abdomens, now confined to southeastern Asia, present anomalous distributions that must be examined in this
connection. 44

The original continuity of the ranges of southern animals, now disOther continuous, is made very probable by such fossil records. in no of in which is fossil evidence examples groups discontinuity
interpreted in the same way. Thus the worm-like the caecilians, are confined to Africa, the Seychelles, amphibians, southern Asia and the East Indies, and tropical America. The South
available

may be

American boid snakes of both Constrictor and Boa have representatives in Madagascar. Many genera of insects are restricted to the southern hemisphere, where their range is discontinuous. The Orthoptera Liturgousa, Stegmatoptera, and Paulctica occur in South America and in Madagascar. The carabid genera Drimostoma and Homalosoma are found in Madagascar, Australia, and New Zealand.

Many

genera of ants are confined to the southern continents.

The

Onychophora, Peripatus and its relatives, are found in Australia and New Zealand, East Indies and southeastern Asia, South Africa, and 4 The ancient earthworm tropical America, including the West Indies.

Primitive Southern

Forms

137

genus Notiodrilus, from which the widespread family Acanthodrilidae may be derived, has a genuine relict distribution. It is found in New Zealand and the Chatham and Snares islands; in New Caledonia; in the isolated oases of central and northwestern Australia, in southernmost Africa, with a species possibly in the Cameroons, in southern South America, and finally in the Central American Cordillera. 30
of these groups of soft-bodied animals are quite unsuitable for preservation as fossils and are scarcely to be expected in the palaeonto-

Many

logical record.

Some

of the regional fresh-water animals, as distinguished from uni-

versal forms, are also confined to the southern hemisphere with discontinuous distribution on the various land masses and islands. Their

origin in the northern land mass or dispersal by this indirect route may explain some of these distributions. Difficulties in the way of

land bridges may not have their directed for In streams favorably the spread of aquatic animals. Africa and Syria, the long series of depressions from the Jordan Valley
this explanation exist in the fact that

to the Central African lakes

dispersal.

may once have been a highway for fish The Central American connection between North and

South America, however, does not appear to afford opportunity for
the exchange of aquatic forms. The conditions governing dispersal of these forms are, to be sure, quite different from those of the terThe primary fresh-water animals, such as crustaceans restrial forms.

and non-pulmonate

snails,

presumably came originally from the

sea.

Fresh-water groups now restricted to the southern continents may be derived from marine forms that had become restricted to the southern
oceans, instead of from ancestral fresh-water forms the northern hemisphere.

now

extinct in

The

three surviving genera of lungfishes are Lepidosiren in South

America, Protopterus in Africa, and Neoceratodus in Australia. This is plainly a relict distribution, as the fossil genus Ceratodus had a

wide distribution, throughout the Mesozoic, in Eurasia, Africa, and North America. The temporary and stagnant waters of the tropics and subtropics presented conditions in which a few forms were able to compete with more modern fishes by reason of their ability to breathe air. Protopterus and Lepidosiren burrow into the ground and For the aestivate for seven to nine months during the dry season. to in live the is able same reason Neoceratodus stagnant waters of pools left by the general drying up of the streams in which it lives. The cichlid and characinid fishes of tropical America and Africa (the cichlids in southern India as well) may have been marine and con-

138

Historical

Zoogeography

fined to tropical and subtropical seas, whence they entered the fresh waters. Their disappearance in the sea may be laid to the rise of the
fishes, which were at first unable to follow into the fresh Such an explanation of the distribution of Galaxias, confined mainly to the extreme south of South America, Africa, Australia, and New Zealand (Fig. 11), is highly probable, since this genus is not

spiny-rayed

waters.

entirely restricted to fresh water.

Members

of the Gobiidae, Cot-

tidae, Syngnathidae, Blenniidae, and Elopidae have entered fresh waters independently in different parts of the world. Thus the five
160

140

120

100

80

60

40

20

20

40

60

80

100

120

140

160

180

20

160

140

120

100

80

60

40

20

40

60

80

100

120

140

160

180

FIG. 11.

Distribution of Galaxias.

now known from New

After Boulenger. A species of Caledonia.

tliis

genus

is

species of the elopid genus Mcgalops are found on the coasts of India and Africa and on the east coast of South America; M, thrissoidcs

occurs in the Magdalena system and in the rivers of West Africa. 10 reviews the distribution of fresh- water fishes and of Darlington

cold-blooded terrestrial vertebrates

in

a thoughtful paper in

1948.

The gastropod genus Potamides has probably been independently

de-

veloped from separate and distinct stocks of the marine Cerithium that have entered brackish water in different parts of the world (cf.

Chapter 12).

The ancient crustaceans of the family Anaspididae, restricted to the fresh waters of Australia and Tasmania and to underground
springs in
tribution.
6 Europe (Bathynella), have a

strikingly discontinuous dis-

Their Paleozoic relatives Uronectes and Paleocaris indicate

a formerly
It is

much wider distribution. no means a mere assumption that the northern forms are by more advanced. Their superiority has been shown by the practical
Eurasian forms introduced into Australia

experiment of introduction.

Primitive Southern

Forms

139

and New Zealand have frequently been able to displace the native forms of similar habits. Thus the two predatory marsupials, the Tasmanian wolf Thylacinus and the Tasmanian devil Sarcophilus,
have survived only in Tasmania, and doubtless became extinct in Australia on account of the competition of the placental dingo ( Canis dingo), an immigrant or introduction from the north, which has not reached Tasmania.

The native Australian songbirds are being crowded out by the forms introduced from Europe, such as the sparrow, starling, blackbird, goldfinch, and greenfinch. The skylark is rated the second worst bird pest in New Zealand. 58 " The goldfish (Carassius auratus), which
was introduced in Madagascar, is displacing the native fishes wherever comes into competition with them. 3 European ants and earthworms have spread in South Africa, South America, Australia, and New
it

Zealand at the expense of the native species. In the reverse direction, of 490 exotic species of animals known to have been brought into Hamburg by shipping in the course of three years, only 5% could maintain themselves, and these only in greenhouses, in tanbark, and in warehouses, not a single species having spread so as to have come into competition with native European forms.- 4 Another example of this phenomenon is found in the mammals that have crossed the Central American land bridge in recent times. The northern emiAmerica have been into better able to South expand and surgrants
vive in the

new

area than have the southern emigrants into North

America.

49

All these considerations support the theory that the forms now inhabiting the southern hemisphere, with a limited or discontinuous distribution, once inhabited the northern land masses, where they then enjoyed the same continuity of distribution as the more modern north-

ern groups, which have driven their predecessors to the south. 27 It has been suggested also that the Eurasian forms are similarly
superior to the North American, and this appears to be true to a certain extent; witness the sparrow and starling, the house rats and

house mouse, the carp, and the host of insect pests introduced from Europe. In the contrary direction, however, certain North American animals have proved themselves almost equally able to spread in competition with European forms, notably Phylloxera (the grape-root
louse)
is

and the muskrat, so that the superiority of Eurasian forms
In so far as such a superiority
it

certainly not without exception.

exists,

may be caused by
Absence of

larger fields.

more stringent natural selection in the parasites and of specially adapted predators
the

140

Historical

Zoogeography

contributes to the success of introduced animals and
to distinguish

may be

difficult

from "innate superiority."

Faunal regions

The

of animals

explanation of the principal features of the present distribution is to be found in the changes in faunal barriers in the

course of geologic time. The study of the actual data of distribution can thus become fruitful only if it is based on the phylogenctic relations of the animals in the various regions and takes into consideration the geologic and palaeontologic data. Faunal lists and statistics, unless subjected to phylogenetic and geologic analysis, are fruitless as a means of inquiry. Regions may thus be distinguished in which

the fauna, at least with respect to certain classes, is more or less homogeneous. The earth was divided into such regions on the basis of its bird faunae by Sclater; and Wallace, principally considering the mammals, has adopted a quite similar division. Wallace believed that this
division into "regions" would apply to all groups of animals, and under the weight of his prestige the delimitation and subdivision of the faunal regions have long constituted one of the principal branches

of zoogeographic inquiry. The belief that there is a division into faunal regions of general validity for all the classes of animals cannot be maintained in the

modern knowledge. For the Mantidae, there is no sharp between the Ethiopian and Oriental regions, which have numerous types in common. The few palaearctic forms are directly
light of

division

derived from these, so that in general, this family can be divided 4 only into two regional groups, a palaeotropical and a neotropical."

Nor can the

distribution of the

non-European genera of mites be

01 brought into harmony with the usual regional divisions. ence to the relations of its earthworms and termites,

With

refer-

New

Guinea

would be grouped with southeastern
affinities

Asia,

whereas

its

other faunal

are primarily Australian. Ceylon, by contrast, agrees closely with Australia in its earthworm fauna. 41 Ceylon termites are clearly related to those of India proper but are usually specifically distinct.*

Chile differs from the rest of South America in

its

mollusks,
is

its

fresh-

water

fishes,

and

its

earthworms, but

its

vertebrate fauna

typically

South American.
exact delimitation of the regions is also impossible, and opinions The* as to their proper limits have accordingly varied exceedingly.
*

An

A. E. Emerson, personal communication.

Faunal Regions

141

boundary between the Oriental and Australian regions has been a special bone of contention, and Celebes has been placed now with one and now with the other, according to the group of animals employed
as a criterion. The creation of transition areas, equally allied to the regions between which they lie, shows that the supposed regions are not objectively defined areas. They are abstractions, combinations of

tent

more or less allied faunal elements, and they will be of varying exand limits according to the animals emphasized.- 8 38 50 Most
'

'

of the major regions is composed of all, faunal elements of quite unequal age and of diverse origin. Thus we speak correctly of a holarctic element in the South American 29 fauna, or of a neotropical element in that of North America. Mayr

important of

the animal

life

remarks on the difficulty of defining the regions, and prefers the term fauna, defined in terms of its origin. Nevertheless, the continuing usefulness of the terms neotropical,
palaearctic, etc., indicates that there is some merit in the geographic partition of the earth as to the main features of the regional animal

The classical major regions of Sclater, and of life of the present day. Wallace 00 were: Palaearctic, Nearctic, Oriental, Ethiopian, Neotrop10 It was pointed out by Heilprin that these ical, and Australian.
unequal value as to degrees of distinctness of and that the Nearctic and Palaearctic, at least, should be combined as a single region, the Holarctic. The argument was carried to a logical conclusion by Huxley, 21 who, employing essentially the same reasoning as Mayr as to origin, proposes grouping the original six regions into the more nearly equivalent Arctogaea, Neogaea, and Notogaea:
divisions are of very
their

animal

life,

Palaearctic
4
,

1

.

..

Nearctic
Oriental

f
J

Holarctic

Arctogaea

Ethiopian (with Malagasy)
Neogaea,

Notogaea

Neotropical Australian

their subdivision into

This matter, with the questions of the limits of the major regions and major and minor faunal or biotic provinces

tive

should receive further discussion in an expanded treatise on descripand historical animal geography.
It is

true that there are

common

features in the distribution of

many

different terrestrial

barriers

and

conditions of dispersal and the the same for many groups. available were highways
animals.

The

142

Historical

Zoogeography

The

differences probably chiefly result from the relative ages of the different groups of animals. Classes that arose at the end of the
Triassic could not spread by means of land connections that were available for the early Mesozoic scorpions, insects, and mollusks.

The
its

older a group of animals, the

more manifold

will

have been

The differences in vagility must be opportunities for dispersal. taken into consideration. Flying animals, such as birds, bats, and
insects,

have been able to spread by routes closed to other land animals, for, even though they do not fly across great stretches of ocean of their own free will, chains of islands and archipelagoes have facilitated their dispersal to islands that were permanently inaccessible
to other forms.

some

When

a land connection

is

of short duration, rapidity

becomes an important factor. Of twenty-two species of amphibians and reptiles on the neighboring continent, thirteen reached
of spread

the British Isles after the glacial period, but only five of these reached Other factors have Ireland, which was cut off at an earlier date.

contributed to inequalities in dispersal. When a new land bridge is established, the emigration of animals from the connected regions will

begin

in

both directions.

Newcomers

in

each region come into com-

petition with the resident forms, and some will be able to maintain themselves through adaptability and superiority of organization, while others will be unable to get a foothold. As a rule, a land connection

mutual exchange of forms rather than to a one-sided invasion of one region from the other. The emigrations of the South and North American faunae via the Panama bridge supply the classic
will lead to a

example of

this

phenomenon.

The distribution of many groups will have the same limits, especially when there are effective barriers. New Zealand, Madagascar, and the Hawaiian Islands are well isolated. Even in such places, the
different groups of animals are not differentiated from their relatives in other regions to an equal degree. Since their isolation, the groups
in the separated regions

may have evolved

in different

ways and

at

different rates.

On

the other hand, non-plastic groups, like

the invertebrates, may have undergone little change. of Madagascar, for example, for the most part represent
ilies,

many of The mammals

whereas among amphibians,

reptiles,

and

insects,

endemic famnumerous genAustralia
is

era are the

same

as those of the African

mainland.
its

highly peculiar in its

mammalian

fauna, while

lizards, butterflies,

and earthworms belong with those of the Oriental region.

The

which the animal

faunal regions are therefore divisions of the earth's surface in life bears a somewhat uniform aspect and differs

Bibliography
from that of the neighboring regions
in

143

consequence of independent evolution during longer or shorter periods of isolation. It is one of
the problems of /oogeography to study the history of the colonization of the several regions and their subdivisions. In the course of this in-

be found that the present fauna of an area is heterogeneous both as to the age and the origin of its various components. The course and direction of the dispersal of the various groups and their order in time must be discovered. These relations may be
quiry
it

will usually

very different for different parts of the same region.
Celebes and Ceylon both belong to the so-called Oriental Region, according But how totally different has been their history! Celebes
a geologically

to the accepted scheme.
is

young

island,

which has received the bulk of

its

fauna over

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.

and

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its

to the geologically

younger mammals.

Celebes,

by

contrast, received

fauna at a time when the

mammals
is

too

had reached a

high stage of development, so that in this island there
41 history of the different groups of animals.

no difference in the

Historical zoogeography

is still

at the beginning of

its

labors.

Much

'

detailed investigation remains to be made, and the insufficient scope of the preliminary work now available makes it difficult to attain a

general view or a single interpretation
future,
as

(cf.

32 Newbigin, 1948 ).

In the

more and more paleontological data become sounder and more illuminating conclusions may be drawn.

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786, 420

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of J. B., et al, 1943, "A discussion of the biogeographic division the Indo-Australian Archipelago, with criticism of the Wallace and Weber lines and of other dividing lines and with an attempt to obtain uniformity in

the

names used

for the divisions."

Proc. Linnean Soc. London, 155:120-165.

51. Simpson, G.

G.,

1940, "Review of the mammal-bearing Tertiary of South

America."

Proc.

Am.

Philos. Soc.,

83:649-709.

146
52.

Historical

Zoogeography
J.

Idem, 1940, "Mammals and land bridges."

Wash. Acad. Sd., 43:137-163.

53. Idem, 1943, "Mammals and the nature of continents." Am. J. Sci., 241:1-31. 54. Stoll, Otto, 1892-1893, "Zur Zoogeographie der landbewohm>ndcn Wirbcl-

losen."

Vierteljahrsschr.
(p. 263).

naturforsch.

Ges. Zurich., 37:233-273;

38:37-77,

294-345
55. Suess,

5

vols., illus.,

Eduard, 1904-1924, The face of the earth. maps.

Oxford, Clarendon Press:

56. Szonto, Istvan, 1940, Forestry, climate,

and drainage in relation to the Htmgarian Great Plain (in Hungarian, with German, French, and English summaries). Roltig, Romevalter: 1-252, 15 figs.
curlew."

P. A., 1942, "The distribution and migration of the hudsonian Wilson Bull, 54:3-11. 58. Taylor, J. W., 1912, "Geographical distribution and dominance in relation to evolution and phylogeny." Trans. 2nd Entomol. Congr., Oxford: 271-294, 5 pis. (pp. 271,275).

57. Taverner,

58. Thomson, G. M., 1922, The
Zealand.
59. Uvarov, B.

Cambridge, Cambridge Univ. Press:

naturalization of plants x, 607.

and animals

in

New

P., 1935, "Ecological and biogcographical relations of Ercmiaii Acrididae," pp. 231-273 in La vie de la region desertique nord-tropical dc lancien monde. Paris, Lechevalier.

60. Wallace, A. R., 1876,

The geographical

distribution of animals with a study

of the relations of living and extinct faunas as elucidating the past changes London, Macmillan: 2 vols., illus., maps. of the earth's surface. 61. Waterschoot van der Gracht, W. A. J. M., et al, 1928, The theory of conti-

nental drift. Tulsa, Amer. Soc. Petroleum Geologists: x, 240, illus. Weber, Max, 1902, "Der indo-australische Archipel und die Gcschichte seiner Tierwelt." Verhandl Ges. D. naturforsch. Artzte, 74:51-62. 63. Weber, Max, and Othenio Abel, 1928, "Die Sa'ugetierc Einfiihrung in die Anatomic und Systematik der recenten und fossilen Mammalia." 2nd cd., vol. 2, Jena, Fischer: xxiv, 898, 573 figs. (pp. 262-263). 63. Wegener, Alfred, 1924, The origin of continents and oceans. Transl., Londoii, Methuen: xx, 212, 44 figs.
62. 64.

Werner, Franz, 1909, "Bemerkungen iiber die geographischc Verbreitung der Mantodeen (Fanghcuschrecken)." Verhandl. zool. but. Ges. Wien, 59: (70)(81).

65.

W etmore,
r

Alexander, 1934,

"A systematic

classification

for the birds of the

world, revised 13:1-11.
66.

and amended."

Smithsonian

Misc.

Collections,

89,

No.

Zimmerman, E.
lulu,

C., 1948, Insects of Hawaii, vol. Univ. of Hawaii Press: xx, 206, 52 figs.

1,

Introduction.

Hono-

67. Zittel, K. A. von, 1932,
xvii,

Textbook of paleontology,

vol. 2.

London, Macmillan:

464, 533

figs.

(p. 332).

O.

The Influence of Extent of Range

THE AREA INHABITED BY A SPECIES (OR ANY PHYLETIC UNIT) IS TERMED The range of an animal is delimited by the lines connecting its range. the outermost localities at which it is found. A species need not occupy the whole extent of its range; for the most part it will be found
only in special habitats that fulfill specific conditions. The extent of a range depends quite as much upon the presence of similar habitats

and upon the arrangement of barriers as upon the history and character of the species. The ranges of animals of different groups are highly unlike. The extent or limitation of the range of a species may
have an important effect upon the selection and constitution of its individuals. Such regularities as are discoverable in these factors are
the subject of this chapter.
subspecies, species, genus, family, etc., that inhabits a small range may be referred to as stenotopic, one with a wide range as

A

extent of a range depends upon a number of factors, including especially the geologic age and variability, the vagility, and the ecological valence of the group concerned. The existing barriers,
curytopic.
of course, prescribe a maximum to the range of any group. of frog on an oceanic island or a carabid beetle in a cave

The

A
is

species

usually

limited in range to island or cave. The range, however, may be restricted to a special part of the island or cave by other factors. The range of an individual species is not fixed but fluctuates with

the passage of time, increases, diminishes, or may be shifted as a whole. The boarfish Capros aper of the Atlantic and Mediterranean was formerly rare on the coast of England but about 1888 became so

numbers were troublesome to fishermen. The moncta was unknown in England before 1890, but since then it has become one of the common English species. Merops apiaster, a bird belonging to the family of bee eaters, has
abundant that
its

noctuid moth

Plitsia

147

148

The Influence

of Extent of

Range

Germany and has failed to become estabits because lished only conspicuous appearance invites persecution on the part of man. The cardinal Cardinalis cardinalis is at present acfrequently entered south
tively extending its range in the Mississippi valley; and the prairie horned lark Otocoris alpestris praticola, a plains species, reached the

The first decades of the nineteenth century. in of the rat the of the extension past century has range Norway great had the effect of reducing the range of the black rat. The limiting factors, on different parts of the border of the geographic range, may
Atlantic seaboard in the

be entirely

different.

capacity for active dispersal or passive transport (vagility) is an important factor in the range of a form. A great range is common among marine animals. Powerful swimmers, such as the tuna and

The

other Scombridae

among

fishes or the

toothed whales
distribution.

among marine

mammals, frequently have a world-wide
trial

Among

terres-

usually the flying forms that are widely distributed. Of genera and families with a world-wide distribution, beetles and butterflies have the greatest number among arthropods, and birds and

animals

it is

bats

among

vertebrates.

fined to birds

among

Species with a world-wide range are conland vertebrates, and principally to butterflies
thistle butterfly

among

invertebrates.

The

Vanessa cardui, which has

such a range, is often seen emigrating in swarms, and such emigrations often invade areas where the species is unable to maintain itself,
as in the British Isles. 41

type,

The milkweed butterfly is a wanderer of this and has spread from North America to the East Indies at a

It arrived in the Tonga Islands in 1863, in relatively recent date. Australia in 1871, and in Celebes in 1873; it has reached the Asores in

the opposite direction and is occasional in the British Isles.'10 The sphinx Celerio lineata, another powerful flier, has spread to all parts of the world. A small range is connected with lessened vagility, when
related forms are compared. The Satyridae and Zygaenidae, with a weak flight, are usually very local in their distribution, often confined
to special habitats or to specific localities.

Such a

distribution

is

un-

known among

the good fliers, such as the Pieridae or Nymphalidae. 38 Flightless birds and birds of weak flight have relatively small ranges, unless they are powerful runners, like the ostrich, or powerful swim28 mers, like the Adelie penguin.

to

may conduce an equally wide range. The inhabitants of transitory bodies of water and of moss may be transported while in their resting stages by the feet of birds or by winds, and are frequently world-wide in
facility

The

with which some animals are transported

The Influence
distribution.

of Extent of

Range

149

This

is

true, for

example, of numerous rhizopods and

other protozoans; of threadworms, one of which, Bunonema richtersi, occurs in the Black Forest, in Switzerland, in England, in the Canary

and on Kerguelen and Possession Islands; of Entomostraca 13 and the salt-crustacean Artemia salina; and of tardigrades such as Milnesium tardigradum. 32 Conversely, only animals capable of active
Islands,

or passive dispersal can inhabit such bodies of water or masses of moss.

The animals

train of civilized

that have been spread to all parts of the earth in the man are further examples of more or less accidental

transport. The Norway rat, house mouse, and domestic dog among mammals; the English sparrow among birds; the flea, various meal worms, various ants (e.g., Camponotus rubripes), and the house fly among insects; the slug Limax flavus and the edible snail Helix aspersa among mollusks; and Eisenia foetida and Helodrilus caliginosus

among earthworms The geologic age
tent of
its

20

may be

cited as examples.

range.

of a group has an important bearing on the exHighways for dispersal available for older forms

may have disappeared before the rise of younger groups. The families of invertebrates, probably on this account, have wider ranges than those of vertebrates. The wide distribution of so many genera of scorpions, pedipalps, and centipedes is doubtless connected with their Among fresh-water antiquity and their now diminished variability. mollusks, the earliest known genera are likewise the most widely distributed, as
ids,

unionids,

and some

by the planorbids, physids, lymnaeids, ancylsphaeriids, all of which are present in the Jurassic as early as the Carboniferous. Anodonta, which does not
is

illustrated

and

appear until the early Tertiary, has a much more restricted range. Greater geologic age seems to be as important a factor in wide distribution of the higher categories, such as genera and families, as

The moths and butterflies are unquestionably more vagile than the beetles; some of them are pronounced wanderers as we have just seen; a few have a world-wide range. This is a rare phenomenon
vagility.

among the species of beetles, but it is a striking fact that nearly all the families of beetles have a world-wide distribution, whereas this is
not true of the majority of the families of Lepidoptera. Coleoptera are an older group, abundantly represented in the earliest stages of the Mesozoic, whereas Lepidoptera are unknown until Mid-Jurassic.
Willis
51

attempts to establish an invariable connection between extent

of range and geologic age of species, but is forced to admit that this relation may be greatly veiled by the presence of physical and eco-

150
logical barriers,

The Influence
by the action

of Extent of
of

Range
factors,

man, and by other

even

among

acteristic of

the slow-spreading plants. A restricted range may be chargroups at their decline as well as at their inception, as is

illustrated

by Sphenodon of

New Zealand

and by horseshoe crabs now

confined to the east coast of North America and to the Moluccas.
extent of the range of a given group of animals is also dependent upon its "ecological valence." Adaptability of any sort favors

The

the establishment of a species in new districts and hence favors the extension of its range. This relation is especially important for terrestrial

animals on account of the wide variety of habitat conditions

on land.
Artemia

The widely transported
and frequently
salina.

inhabitants of temporary ponds

are eurythermal

The

also euryhaline, like the salt-crustacean resistant nature of Cyclops fimbriatus is shown

by the variety of situations that it inhabits. It is found in Greenland and Ceylon, on the plains as well as on Mount St. Bernard. It occurs
in concentrated mineral waters, in iron-ochre deposits of brooks,
in caves

and

and mines. 46

Euryphagy is frequent in widely distributed forms. The food of carnivorous animals is least restricted, since mammals, birds, and
fishes,

and even

insects,

worms, and mollusks, are much more uniform
fruits, seeds,

in

chemical composition than are leaves,

plants.

Thus the most widely

distributed

mammals

or other parts of next to the bats

and marine forms are the carnivores such as the Eurasian wolf, with the closely related North American wolf; the leopard, ranging through Africa and south Asia into China and Borneo; or the puma, whose range extends from Alaska to Patagonia. Both nocturnal and diurnal birds of prey may have a very wide range so the sea eagle, the peregrine falcon, and the barn owl. The extraordinary euryphagy of the raven, which feeds on carrion and on living animals from roe fawn and birds to insects, worms, fishes, birds' eggs, plants, and seeds, must have had an important effect in giving it its wide distribution.

There

is

of a species

a relation, though sometimes obscure, between wide range and position at the top of a food pyramid and between

wide range of the individuals of a species and that of the species as a whole. A larger carnivore, in general, will have a wider range
than will the smaller herbivores that constitute
its

food.

Euryphagy has plainly furthered the spread of herbivorous insects. The caterpillar of the nearly circumglobal cotton bollworm Heliothis
armigera feeds mainly upon the tobacco plant at Delphi, on maize in

Restriction of

Range

151

23 The caterpillar of Java, and on the cotton plant in North America. the widespread Utetheisa pulchella is no less catholic in its food habits

and

ences. 37

also displays a high degree of tolerance toward climatic differHelix aspersa, transported by man to all parts of the earth,

has unquestionably been able to establish itself on account of its 45 Higher animals may show great latitude in foods eaten. euryphagy.

The mourning dove
deer 516 species. 3

takes 149 species of plant food, 33 the white-tailed This helps to account for the wide distribution of these animals and of related forms. On the other hand, animals limited to restricted environmental niches where food is scarce, as in

mountain

torrents, tend strongly

toward euryphagy.

Restriction of range

Limited range, especially for species and genera, is, on the whole, a much more general phenomenon than wide distribution. Specialization,
i.e.,

affords so

the exact inherited adaptation to given habitat conditions, many immediate advantages that specialized species usually

win

in the struggle for existence.
lies

The advantage

of flexibility of or-

ganization

primarily in the capacity for

wider distribution.

The

fatality in specialization lies in the corresponding loss of the capacity

for adaptation.

appear only undergo decided changes, which tend to unable to meet these particular changes.

The disadvantages of adaptation to a special habitat when the habitat conditions in the range of an animal
kill

off specialized

forms

The

limitation of range of a species

may be

extreme.

Small islands,

high mountain ranges or peaks, mountain valleys, and other sharply defined areas may have species confined to them. Examples among Lepidoptera are to be seen in the Attacus isabellae, known only from
a forest near Madrid; the notodontid Rhymatophila alpina, from the neighborhood of Digne in southern France; the

known only hawk moth

Akbesia davidis, from Akbes in Syria; and the genus Zygaena, with nearly 200 closely similar forms, most of them in the Mediterranean
district

and many with very small ranges.
is

The

flight

area of Z.

scriziata in Algeria

sometimes restricted to an area 8 to 10 m.

broad. 39

carabid beetles are restricted to particular mountain ranges or to parts of them; Carabus adonis is known only from the Parnassus and Taygetus, and C. olympiae only from the val-

Many

flightless

Among snails Radix inleys of Aosta and Sesia in the Apennines. valuta is restricted to a small mountain lake in Ireland, and Papilifera scalaris is found only in a small calcareous area in Malta; one mollusk

4

152
genus Lanzia
is

The Influence

of Extent of

Range

confined to the moss of a mountain peak in Bourbon. 8
is

Among

birds a restricted distribution

common among humming-

birds, as in Oreotrochilus

chimborazo chimborazo from Chimborazo In and Eriocnemis glaucopoides from Valle Grande in Bolivia.

seaside sparrow is a full species. It inhabits a coastal grassland area about six miles long and no more than a half-mile wide. No other suitable habitat for this bird has been reFlorida, the

Cape Sable

19 ported for a long distance on either coast of Florida. The combination of slight vagility with the presence of barriers to

It seems likely that stenodistribution usually produces stenotopy. topic forms will be found to have in common other characteristics

that influence

limited distribution

the limitation of their range. Various reasons for be considered: slight power of expansion, may

high degree of specialization in adaptation to special conditions, youth of the species, or age and decline. Low vagility alone does not require consideration on account of the
available.

amount

of time that has been

Germany covered with ice during the glacial have been period completely repopulated by such slow-moving forms earthworms. The power of expansion possessed by a snails and as
parts of

The

species

must be associated with

its

fecundity, but no example of re-

stricted range certainly assignable to a low degree of fecundity can be cited. forms, in general, will arise in quite definitely circumscribed areas, as is to be seen in the melanistic forms of the

New

Lepidoptera Boarmia consonaria and B. consortaria in the neighborhood of Maidstone in England. 17

The melanic

variety, carbonaria, of Biston betularia, the

peppered

moth, which appeared near Manchester, England, in 1850, has completely superseded the non-melanic parent species in parts of England

and France.
This phenomenon, referred to as "industrial melanism/' has successively attracted the attention of butterfly collectors, ecologists, and
geneticists.
is

so

much

It appears that the dark form is a recessive mutation that favored by its protective resemblance on smoke-blackened

walls, wrth a differential exposure to predation of the light-colored dominant form, that it rapidly increases in abundance wherever such

backgrounds appear.

reviewed by Ford. 15 The fish Atherina riqueti* which has been in existence only since the end of the seventeenth century, has appeared in the Canal du Midi between Garonne and the Mediterranean. It is a fresh-water derivative of the marine A. boyeri. The surviving relicts of ancient

The matter
4

is

Extended Ranges
forms frequently have a restricted distribution.

153

Thus the Australian

hmgfish Neoceratodus forsteri is known only from the Burnett and Mary rivers in Queensland, whereas its extinct relatives were widely
distributed; the ancient rhynchocephalian

Sphenodon

is

confined to

New Zealand; the duckbill Ornithorhynchus is found only in the streams of southern Australia and Tasmania; the genus of snails
Pleurotomaria

now

occurs only in the Moluccas, Japan, South Africa,

Here the very barriers that limit the distribution of the forms in question have conditioned their survival by keeping out more efficient competitors. It must be added that a small range is not
and the
Antilles.

The singular cephaloone besides with the Nautilus a chambered shell, only pod Spirula, and Indian oceans. The genus the Atlantic, Pacific, ranges through
at all a necessary characteristic of a relict form.

Eukoenenia, a small primitive arachnid of the order Palpigradi, has been found (represented by various species) in the Mediterranean

and Paraguay. Local species arise among land birds on islands, in spite of their powers of flight. Such endemism may result on account of their very vagility, which, combined with a homing instinct, may condition the
region, in Texas, Siam,

return of wandering individuals and thus increase the effectiveness of
their insular isolation.
30

In contrast with the relict forms, the wide distribution of a group combined with the appearance of numerous differentiations, such as numerous subspecies in a single species or of numerous species of the same genus, is an indication of vitality. Widely distributed genera tend to have numerous species. A world-wide (cosmopolitan) range is the maximum possible, and it stands in contrast with geographic
limitation.

Extended ranges

An

animal

is

said to

be cosmopolitan when

it

occurs in

all

the

places that afford it suitable habitat conditions; cosmopolitanism does not mean that it occurs alike in salt and fresh water and on land.

found to inhabit a wide variety of unlike habitats The term cosmopolitan distribution is used is said to be ubiquitous. in the geographic sense, and ubiquitous occurrence in the ecologic. Cosmopolitan species may accordingly be marine animals occurring
that
is

An animal

in all oceans or air-breathers occurring in all the

zoogeographic regions, although the polar regions are often omitted from consideration in this connection. The widest distribution in both the geographic

154

The Influence
among

of Extent of

Range

is probably to be found among Protozoa, espethe rhizopods. Thus Cyphoderia ampulla (Fig. 12) lives in the ocean, in salt-water marshes, in fresh water, in the sand of the

and ecologic sense

cially

warm ponds
protozoan
is

seashore, in the springs and glacial lakes of mountains, and in the of the lowlands. In addition to central Europe, this

sian Lapland,

recorded from the coast of the Arctic Ocean, from Rusfrom the Rocky Mountains, and from Argentina and

FIG. 12.

Cyphoderia ampulla.

After Scluilze.

The thistle butterfly Vanessa cardui is an example of a metazoan. It is known from all parts of the world distributed widely with the exception of some small islands and South America, and
Paraguay.
ranges from the tropics to the arctic regions and from the plains to the snow line in the mountains. Animals found in all seas are the
edible mussel Mytilus edulis and the thresher shark Alopias vulpes. World-wide distribution within the tropics has been referred to as
circumtropical. Such a distribution is characteristic of many of the families and orders of animals. The geckoes among the lizard family

Gekkonidae, and the order Crocodilia, the alligators and crocodiles,

more common for higher than for lower taxowide categories. range is much commoner for genera than for species; it is even more common for families and orders. The families of myriapods and scorpions, and very many families of inFresh-water fishes, amphibians, and reptiles sects, are cosmopolitan. have no cosmopolitan families, though Siluridae, Ranidae, and Coluis

afford examples. Wide distribution

nomic

A

bridae are very widely distributed.

Numerous world-wide genera

of

air-breathing vertebrates are to be found among the birds; witness Turdus, Hirundo, many birds of prey, owls, and numerous water birds. Canis and a few genera of bats are the only genera of mammals that

Extended Ranges
closely

155

approach such a range. Species of such wide distribution are not uncommon among the marine invertebrates. Among terrestrial animals there are a few cosmopolitan species of arthropods,* but among vertebrates the only cosmopolitan species are a few birds, such as Pandion haliaetus and Asio accipitrinus. Species with a wide range usually belong to genera with a still wider distribution. 9 Thus the tiger, leopard, and lion belong to the more widespread genus Panthera, and the widely distributed wolves to the almost cosmopolitan genus Canis. The genera to which the raven Corvus corax, the peregrine falcon Falco peregrinus, and the ringdove Columba oenas belong are both widespread and rich in The same may be observed among reptile and amphibian species. genera (Lacerta, Bufo, Rana) and among invertebrates such as the land and fresh- water mollusks (Helicidae, Lymnaeidae). Species of monotypic or oligotypic genera, in contrast, usually have a narrow range. Among American moles the genera with the most restricted distribution are the least varied. 48 Other examples among mammals are the panda (Aelurus), the giant panda (Aeluropus), and the binturong (Arctictis), and among birds two Brazilian species of Eurypyga, Mesites with a single Madagascan species, and Didunculus with a single Samoan species. 1J Eigenmann found in the course of his researches on South American fishes that in a given river system, even so vast as that of the Amazon, the widespread genera have several times as many species as
the genera confined to the system in question. This indicates that a genus with a small range tends to have fewer species because there is
less available

habitat niche.

space for them, with, of course, less possible variety of The examination of the conditions that make possible

a wide range will cast
especially vital

some

light

on

this

problem.

It is precisely

the

and adaptable genera that are differentiated into numerous species and have a wide range. Such genera are in the minority. Exact statistics show that among the most widely distinct families of plants and animals the monotypic genera are by far the most numerous, followed by those with two species, then by those

with several or many. 51

Many
*

stricted varieties at the borders of their

widely distributed species tend to develop numerous rerange while remaining conrufipes, Tegenaria derhami); (Vanessa cardui, Celerio lineata, Heliothis armigera, Nomophila among Lepidoptera; Cercyon nigriceps among Coleoptera; and

A

few arachnids (Theridium tepidariorum, Th.
insects

a

few

noctuella) 30

the myriapods Scolopendra morsitans

and

S. stibsfyinipes.

compared with 70-80 mm. Portugal. Scotland. Corsica. especially in the somewhat isolated outposts whose inhabitants do not interbreed as freely with the central mass of the species. 21 in the Pyrenees) and of the "Formenkreis" Levantina in Palestine. and west. or they will die of starvation. Alsace. in the southwest and west by P. occurs in central a form somewhat intermediate between the subspecies in the Russia. which varies strongly at both the horizontal and vertical limits of its range (for example. as in H. or each pair of individuals.. Parus pleskii. frequently has albinos in the southern part of its range. Spain. Among spective size. caeruleus and its 7 subspecies. and the separation of the individual stations at which the species occur will be less. which shows unusual variation at its southern limit.156 stant over The Influence of Extent of Range wide areas in the center. It is probable that the conditions of life at the limits of the range of a given species are less favorable and that these are responsible for the change and dwarfing of the species. males of Triturus palmatus from Porto. flavipectus. in Italy. and Algeria. The same is true of Helix (Arianta) arbustorum. north of the Pyrenees/' Another instance of this relation between distribution and variation titmice. presented by the European birds of the genus Parus. The European newts sometimes have constant dwarf races in the south. toward the northeast and north it is replaced is by Parus cyanus. minimum value for the extent of the range. which is rarely albinistic in Germany. south. Albinism is reported by Paul Hesse (personal communication) to be especially common at the limits of the specific range. as well as at the eastern limit in Japan. carnivores must be much fewer than their prey. has a special area of its own within v^hich the presence many . This is true of the snail Helh in Algeria. being dependent upon them. which dethe nature of the species and consequently varies with pends upon the species concerned. 2 measure 55 mm.* The Eurasian deer have formed dwarf races at the western limits of their range in the Faeroe Islands. called The least specialized form. and that a flying insect or an antelope will demand a wider space in which to move than a snail or a sloth of similar re- Carnivores will require a larger area per individual than herbivores. east. and Helix (Cepaea) nemoralis. species of carnivorous animals an individual. and in the east it reaches the range of P. Each individual animal requires a minimum There is a space in which it can satisfy its needs for food and movement. It is evident that an area that would seem large to a mouse would be small for an elephant. In a narrow range the habitat conditions will in general afford less variety. arbustorum in the Alps. aspersa.

and even the centipede Lithobius is and dippers (Cinusually found singly under a given stone. and this dividual pair. moles. 29 quired by vary from The size of suitable territory apparently redifferent species of birds that do not nest in colonies may 49 to about 60 square 0." Different kinds of vertebrates. territorial habit is widely developed among Kilda wren defends from trespass a far larger area is required for its food.Territoriality 157 not tolerated. including particularly fishes. Territorial defense is also known among certain invertebrates including.2 of an acre for the wood thrush The territory held in the breeding miles for the golden eagle. the fiddler crab. however. (7) predators are more easily avoided on account of the close acquaintance of the territory holder with the given locality. territory has been defined as "any returns. (2) limits the breeding population and hence helps control the population size within the limits of carrying capacity. The concept It defended area. clus). in- of territoriality is related to the idea of home range. foxes. Uca. The territorial habit usually has one or more of the following probable values: (1) produces a more uniform occupancy of local frag- ments of the habitats. eagles. (5) with larger territories. (8) apparently for psychological reasons. . kingfishers (Alcedo). (3) furnishes partial protection of helpless young. with the area or even with the spot to which an animal deal may In a more narrow sense. 2 species. among others. Territoriality number. (6) parasites and disease spread less rapidly. in contrast with the herbivorous milliped Julus. there is increased vigor of defense by the occupant and decreased vigor of attask by an intruder of the same birds. birds. provides a reserve of unmated males and females. A of pairs. lizards. and mammals. and ants. tends to ensure an adequate supply of near-by food. of another animal of the same species is This is true of bears. (4) when well developed in territorial birds. actively defend a part of their total range especially during the breeding season. 10 season is smaller in years when the population is more dense. is essential to preserve the species from inbreeding and from "accidental" extermination by adverse con- minimum number ditions. allowing more prompt replacement of a mated animal that dies. The limitation of the number of pairs able to breed in a given area may on occasion be controlled by the pugnacity of the male birds. but this limit is seldom reached in popu- than lations of land birds. 18 The The St. multiplied by the area required by an would express the minimum range of the species.

ditches. though present on the mainland opposite and present on the much smaller island of Bali ( about 5000 sq. and are for the most part much smaller. Thus trout. although the widest known ranges are those lakes of Europe. since available areas are too small to support larger forms. etc. Galba truncatula. and the spring snail Bithynella relicts are relicts of the ice age. are not to be expected on small islands. The absence of coastal species of sharks from inshore waters of the small Pacific islands. extermination of rabbits on Laysan and Lisiansky Islands. 50 The usually range from the size of the hamster down. springs. the Hawaiian Islands for example. Thus the Panamanian caiman is driven from the larger bodies of water by the crocodile. small clam Pisidium. such as the North American "sheepshead" Aplodinotus grunniens. 14 The introduction and selfspace is not sufficient to maintain them. like the cattle that were landed on New Amsterdam 7 No exact definition of the size of islands that can (66 sq. km. and puddles.. The tiger is absent from Ceylon. and the giant It catfish Silurus glanis in the thus appears obvious that the in animals of general require a wider range than do larger species smaller related animals. The smallest bodies of water. such as the musk ox or reindeer. km. and Large animals. may be explainable on the ground that the available . and relicts of the steppes. some xeric snails and grasshoppers are . Perhaps the small ponds proIt is in question vide a refuge from larger enemies or competitors for which the area would be too small. the arapaima Ara- paima gigas larger rivers in the Amazon. Pisidium and leucostoma. the tiger may have been exterminated in Ceylon by man. the Some small mollusks.). described by Wetmore. the turbellarians Planaria alpina and Polycelis cornuta. in consequence. Only small animals have been able to maintain themselves as relicts of the ice age or of the steppe period in central Europe. the larger fresh-water fishes occur only in the larger bodies of water. about 25 kg. Also we may be dealing again with a direct effect of size of habitat upon growth size. ) The size of Ceylon cannot therefore be the determining factor. On the other hand. is an instructive phenomenon in this connection. maintain a given species is possible.158 The Influence of Extent of Range Effects of space limitation very likely that other factors affect the problem of minimum range. harbor only small mollusks such as the spring snail Bithynella. of the protozoans. though if introduced they may be able to maintain themselves for a time. certainly are more frequently found Gyrorbis in small bodies of water than in large.

600. which is much larger than the Varanus salvator of the larger neighboring inlands and the mainland. The vast multiplicity of may not be expected on an island. 200. the specimens grown in 100. km. the physical conditions in these basins may be the determining of animal It is probably true. 9 Iceland has fewer species of birds than Norway. with Limnaea stagnates. to this relation appears to be the Komodo Island monitor Varanus komodoensis.. and 18 mm. has only 181 species of birds. of water attained a shell length of respectively 5. 9. Thus the size of the pelagic copepods of the genus Diaptomus depends on the size of the bodies of water they inhabit. and 2000 cc. Although the causes at the bottom of this relation between size and range are not yet clear. a large number of facts may be brought together from this standpoint. 12. field ferent causal factors involved. 20 Trout from the Aar averaged 240 gm.. . however. than uncircumscribed areas of similar animals that characterizes a continent size. those from its affluents. extent of the space available has a direct effect upon the conExperiments have shown that the individ- uals of a species reach different sizes varying with the amount of 40 made space given them. their size remained smaller than when this The results of numerous recent researches in which confirm and extend those of Semper have been 1 comprehensively reviewed by Allee. Sikkim. Dwarfing in consequence of reduced range may also be seen in the numerous Alpine animals whose ancestors had a much wider range in the glacial times and were much larger. and if larger numbers of specimens were kept in the same container. whereas New South Wales has 385. 132 44 The whitefish Coregonus from the small lakes in Switzerland are gm. 35 Fresh-water mussels Unto and Anodonta have smaller individuals in brooks than the same species in rivers.. In the original experiments of Semper. in the same length of time. Small well-defined situations like islands or oases have fewer species factors. Tasmania. whose bird fauna richest in the world. has 500 to 600 species of birds in is one of the 4015 sq. that the frequent presence of dwarfed forms of animals on small islands depends on space relations. according to Gould's figures. who discusses at length the difthere were fewer. 16 The smaller size of most of the animals of the Baltic and Mediterranean seas compared with those of the Atlantic probably does not fall into this category.Effects of Space Limitation 159 The stitution of its inhabitants. Their formerly continuous range has been reduced and split up by the encroachment of the 43 A conspicuous exception forest into the mountains from the plains. dwarfs compared with those from the Lake of Zurich.

and thereby increased On formation of species by adaptation. Island sq.690 268.461 118. possibilities for the Area. kin. The greater extent of the oceans in the is A somewhat similar exhibited southern hemisphere similarly reacts favorably on the evolution of genuinely pelagic animals. This is evident when the amphibian and reptile faunae of various large islands with similar climatic conditions are compared. and the Cape Verde Islands (3851 palace garden in Bonn. has 43 species of birds nesting within its confines. of about 22 acres. km. has only 337.100 501.* In similar climatic zones the larger stream tends larger to have the larger fish fauna. The Azores (2388 sq. The same * relation appears among the plankton. . A principal reason may be that they afford a wider range of habitat conditions.859 9. The West Indian islands.234 243 181 88 61 100 Java Ceylon New Zealand 200.132 131. Indies shows that the extent of the area concerned when result though with equally necessary exceptions the numbers of species of fishes in the faunae of the by river basins. These seas have larger numbers of whales. apply to parts of a mainland. German 2" Chun 6 characterizes For details.) have fewer than 30 breeding species.833 22 38 37 2 106 157 97 15 Cuba Ilispaniola Jamaica Puerto Rico 77.733 65.563 443. about 45 times as large.314 23 29 16 56 66 28 30 13 The comparison factor only of the New Zealand fauna with that of the East is the deciding other factors are similar or comparable. see the original edition of the present work.) only 38. on islands. Numl>er of Sj>ecies Amphibians 85 Reptiles Borneo Madagascar Sumatra Celebes 715. are factors that do not the other hand. in general have the richer fauna. are comparable only with one another. The lessened possibility of chance and increased difficulties in the way of the immigration of new forms. with other conditions equal. likewise. This difference is necessarily conditioned by the fact that the least sq.253 10. km. 27 A range of a species in an interbreeding community is larger than the area required by a single pair of the species within the range.160 The Influence of Extent of Range while Celebes. broader areas.

. Chi- . of 143 species of decapods in the found on the east coast and only 65 on the west. 1931. and the prosperity of a species.. of Chicago Press: ix. Regions with a wide variation in habitat conditions contrast with such uniform areas. their giant birds. and extended mountain ranges are especially favorafford residence able to the development of specifically adapted forms. and otherwise rich animal life. and these favor the existence of numerous species and subspecies. 116 are while on the east coast former greater extent. as among echinoderms >2 or among pycnogonids. cago.4 The broken and varied east coast of the Adriatic affords a greater variety of habitat than the more uniform west coast. There unquestionably a certain relation between the size of a region and the variety of its fauna. Wallace applies the same reasoning to Australia. steppes. 431. but the littoral species have much more circumscribed ranges than the deep-sea forms. Univ. W. prairies. The subdivision into varied situations of the sea bottom of the Sagami herrings Bay near Tokyo has its share in conditioning the 11 The Norwegian coastal surprising wealth of species in this area. The variety of conditions in the littoral affords habitats for a greater number of gorgonians than the deep sea. a study in general sociology. itself the maximum wealth of species is reached 31 in the broken Dalmatian coast with its islands.Bibliography 161 the Antarctic pelagic fauna as in general richer in comparison with that of the Arctic. show much more variation and division into races than the 42 on account of herrings of the open Atlantic and of the North Sea. with their giant land turtles. This has been postulated for the Seychelles. are in general more favored by wealth of individuals. which have 82 Antarctic species as compared with 62 in the Arctic. Animal aggregations. on this account. Alice. It has been supposed. since they afford conditions adapted to needs of varied animals. even among the bottom-dwelling forms the Antarctic fauna is the richer. BIBLIOGRAPHY 1. its further development and progressive adaptation. C. 6 Large areas with uniform conditions like great stretches of forest. that a richly differentiated fauna in a relatively small area would warrant the supposition of its Adriatic. but it cannot be expressed in figures or otherwise sharply enough defined to afford a sufficient basis for conis clusions of this nature. 35 figs. the great variation of physical conditions on the coast. since they and perpetuation to large numbers of individuals.

Frankfurt. Schmidt. 1 dem arktischen und map (p. L. 1937. "De la dissemination des especes vegetales et animales. 47 ff.. K. H. 195). Teil." 18.. H. 204. C. 2. 1915. 1945. 22. Acad.). 18 pis. "Pycnogonides. xii. 1897. 48 pis. B. 100:1430-1436. 1898. "Ober die Entstehung des neuzeitlichen Melanismus der Schmetterlinge und die Bedeutung der Hamburger Formen fur dessen Ergrunteleuropa. Rud. B. Vogt.." figs. Bouvier. "White-tailed deer foods of the U. Koehler. Alfred E. 32 maps. Cambridge Natural Hist. Applcton: 2 vols. Wiesbaden. 33). Charcot" (abstract). Ekman. Deuxieme expedition antarctique franZentralbl Zool cais (1908-1910) commandee par J. The Influence W. London. Wilhelm. Ford. 20. : 15. Ostasienfahrt Erlebnisse und Beobachtungen eines Naturforschers in China. Darwin. 311 (p.. Blanchard. Game and Fresh Water Fish: xxiv.. Tiergeographie der Selachier.. 11 2 maps.. Jena. Doflein. 4:1-40. 1:392-399. S.. 279). A. Syst. pp. und Ceylon. 1900. (vol. 1895. R. 178. Eigenmann. 10. Condor. 1 fig. The principles of animal ecology. 5:349-351. E.. 16. Die Beziehungen ztvischen Stuttgart. 37:567-600.-n." (p.162 2. Leipzig." /. Origin of species by means of natural selection or American ed. 58 pis. 118. A. C. 3:1-459. 579. Carl. Jahrb. 836. 594. Zentr." . Jordan. Orlando Park.. Kobelt. E.). Alice. Hofer. maps (p. and Karl Philadelphia.. Natl Museum. 11. of Extent of Range P. Engelhardt. Exped.. xxiv. Teubner: xiii. 278). Suppl. 1906. illus. Dixon." Biol. 18). 1885. California.. Y. 1 map (p. Fischer: vi. illus.. 558. ophiures. 3.. Franz.. 8 figs. 2 8. Emerson. Hermann. 6. 1932. 263 figs. pis. and R. 462). Butterflies. "Echinodermes (asteries. Tallahassee. S. Saunders: Wildlife Afwirig.. 1-64. pt. Niigele: tischen Plankton. 1949. Kreidel: x. J. 353). 4. the preservation of favored races in the struggle for life. Die Susswasserfische von Mit- 17. et echinides). 1913. 72 text figs. 368. der Muscheln aus der Familie Najades. 20). Murray: xiv. Lam. (p. 3:1-110. E. Munchen (m. Japan. 1881. sildpol. 1920. Thomas Park. H. London. und B. 4 maps... Akad. 1941.. Atwood. 1914." "13. Florida bird life. "Molluscs. Cooke. 19. Howard. maps (pp. 29:49-56. L.. Charles. Collins: xiv. 390 figs. fimile. Biol. Aus den Tiefen des Weltmeeres. 12. U. Dept.. Wilhelm. N.. dung. Die Mollusken der meridionalen Sub-Region. Pans. C. 5:314-332. Grote. Ergeb. S. 352). Chun. Schilderungen von der Deutschen Tiefsee-Expedition. Studien zur Zoogeographie: 2. "Monographic der Selachier der Miinchener zoologischen Staats-sammlung I. "A catalogue of the freshProc. 7. Howell.. 1909. 46 pis. Ifg. illus. 5. 368. 30 pis. Eigenmann.. "Einfluss des bewegten Wassers auf die Gestaltung life. 21. antark- Idem. 6 E. from 6th English ed. H." 9. Territory in bird figs. 1891. 511. Sven. 5. (p. 1899.. 14:1-81 (p. sci. subantarktischen Binnengewassern. "The golden eagle in San Diego County. Hasebrock. schwed. R. Zool. water fishes of South America. 14. Fla." Abhandl." C.. Wiss. und Copepoden aus antarktischen und "Cladoceren 1905. 1914. 4..

89-100 in Fifty years of progress in American Ornithology. 133:1-10. Y. 1916. sp. 2 maps (p. 13-14. Y. of the birds of the Archipelago. Roule. N. "The theory of territorialism and its development. list C. deut. all maps (p. (p. Nijverheid en Handel: 1-663 (p. Heft 21:1-188. 38. 1902. "Arktische Tardigraden. 37. 400). (vol." Am. "Essay on the zoogeography of the Pacific Islands. "Atherina riqueti nov. vivant dans les eaux donees. "Ueber das Wachsthum von Lymnaea Verhandl phys. Louis. Jr. Pesta. Lohmann. 5-7. 1872. 14 pis. Michaelsen. "Die Appendicularien der Valdivia Expedition. Schmeil. M.. 36. nouvelle espece d'atherine Zool. "Deutschlands freilebende Siisswasser-Copepoden : III. illus.). 15-16. 1895-1899. . a systematic known macrolepidoptera edited with the assistance of wellStuttgart." Semper. with a Hist.. Sharp. 293 figs. 136 ff.. 1896. Friedlander: vi. Deuticke: x. Anz.. Idem. Museum Natural 28. 626. Nice. 1933. Otto. "The marine ornithology of the Cape Verde Bull. Paul. Museum Natural Am. study of the song sparrow. 33.. Idem. 1909. illus. francaise (1908-1910) [Abstract] Zentr. Linnean Soc.." Naturwissenschaften. 2. J. N. 163 commande'e par Charcot. Richters. Macmillan)." pp.. 6:xii. N. . Hist. 20). C. J. Java zoologisch en biologisch. 19-30 (p. Jahrb. Biol. Zool. map (p. "Die systematische Stellung der Zygaeniden. Sidney N. Seitz. Wiirtzburg. 1912. 2 figs. Schmidt..) 3:271-279. Schnee. Syst. 1922. Hans. Murphy. 1918. Idem. 1939. 500. 1911-1915. 39:1-3. Pagenstecher." Verh. 4-256). 34." Fauna Arctica. Dept. 25. Willy. K. pis. Berlin.Bibliography Deuxieme expedition antar clique J. 50:211-278. 30.. 298.. Putnams: xv. 1948.. 4: 657-663. Die Dekapodenfauna der Adria. Ges. Walter. Rundstagnate. Koningsberger. specialists. Die geographische Verbreitung der Schmetter451. Arnold. 181). 186. Landbnuw. 20:387-412 (p." Cambridge Natural Hist. David." Wildlife Research Manag. Rosene. 26. 1924. Zool Ges. 29.. P. (n. Y.. 31. 32. 4 figs. 1930. "Die geographische Verbreitung mariner Bodentiere. "Die Landfauna der Marschall-Inseln nebst einigen Bemerkungen zur Fauna der Inseln." schau. 40.. Buitenzorg.-med. 460 ff.). 2 vols. 25:262-267. Die geographische Verbreitung der OUgochaeten. Leipzig. 9-11. 10 27. Islands." pp. Adalbert. Oceanic birds of South America." Zool. 10 figs. 24. Bibl Zool.f. 291). 36 ff. (repr. 23. Fischer: 1 ix. Jena. Leafl.S. Entomol. "Studies in the life-history of the song sparrow. 1903. "A preliminary investigation of the food habits of the mourning dove in Alabama. M. 275-292 in Shurcliff. 353).. pis. 3:493- 508. [incompl]. 1933. pp. (p.). 5: 279-280. 4:1-243. 1904.. 1904." illus. W. 1906 account of all the known 39. Carl. 1914. The macrolepidoptera of the world. Lehmann: 14 vols. B. Ferdinand. Jungle Islands.. 1936." 35. "Insects. Otto. Kiikenthal. Centropagidae.. A population 290. R." Trans. 41. linge. 24:157-192....

form des Hermelins Putorius ermineus (L. Proc. G. 48. Willis. Wetmore. 47.). Brit. 49. Willy. W. Wolterstorff.. naturforsch. Land and freshwater Mollusca of Taylor. 260-266. 1896... Wochschr. 1913.. pts." Naturw. 6 figs..f. Storch. 1914." 6th Intern. 46 text figs. F. "Ueber die Ergebnisse der ersten Laichfischfang-Statistik des Kantons Bern pro 1913-14. in the life history of the wood thrush. Thienemann. Natl True.. 48:76-108.. Tristram. "Ueber Putorius ermineus minimus Cavazza. 10:16-23." 51. 788 ff. the British Isles. 248. Hydrobiol." Museum. W. Congr.. Natl. J. species.. "Zwergformen der palaarktischen Urodelen. "A revision of American moles. Adv. "Die Factoren. U. xvii. Weaver.). life among lava rock and coral sand: the chronicle of a scientific expedition to little-known islands of Hawaii. Surbeck.. Cambridge Univ. S. 44.. 46. Fischerei Zeitung. 1925. Berne: 258-263. H. F. 16). Assoc.164 The Influence (n. 1904. Alexander. 4 pis. 1910-11. 1939. illus. 259. 1893. (p. Th. Presidential address. Bern. 1922. 45." Schweiz. 1913. G. 43. Rept. Press: vii. 2:225-368 (pp. "Die modernen Heringsforschungen. (p. B:625-631. 52.). Ges. J. xviii.. illus. 22:228-238. Set. 8:269-288. 1 pi." 50..) of Extent of Range 42." Arch. "Bird Geographic Mag. 1914. a study in geographical distribution of C. 1913:79-91. Otto. 19:1-112. 332). "Studies Bird-banding. 1893:784-797 (p. B. Ein ZwergStuder. C. Cambridge. 90 ff. Zool. welche die Verbreitung der Siisswasserorganismen regeln. Age and area. ." Mitt. August. R.

The whole space 13 occupied by living organisms has been called the "biosphere/' The biosphere presents extremely varied aspects in its various parts. From the botanical viewpoint we already have a completely worked-out classification and nomenclature for the major parts of the biosphere. study of the biosphere thus requires a subdivision into the component parts corresponding to these differences. It is true that the zoological subdivisions will frequently be based upon the botanical. The primary topographic unit is the niche or biotope. with their consequently completely different mode of distribution. such as forest and grassland. Furthermore. Such a unit is an area showing uniformity in the principal habitat conditions. ocean. and on account of the relatively greater importance of marine and fresh-water animal life. are themselves in turn the habitats in which animal life is found. whereas plant geographers have long given serious attention to biogeographic studies upon an ecologic background. Little has been A done in this direction by zoologists.the including both land and sea. 2 5 ' Zoology cannot take over a phytogeographic system without reOther principles of subdivision must frequently be applied vision. but not for long periods. but It rises it penetrates only a few meters deep in the solid earth. but they must not slavishly follow the botanical scheme and must find new principles of their own as the occasion demands. The biotope is as basic for the ecologic subdivision of the biosphere as the species is in the systematic classification of living beings. what are primary divisions for the botanist. Biotopes and Biocoenoses ANIMAL AND PLANT LIFE OCCUPIES THE WHOLE SURFACE OF THE EARTH. though 165 .C7. Life extends into the depths of . although unfortunately no unanimity has been attained in this respect. both according to the habitat conditions and the forms of life represented. higher into the air. for animals on account of their vagility.

zones. hills." Among the biochores differences in general aspect. the so-called "habitus. The biochores and superbiochores are finally combined into still higher groupings. and high mountains. Thus. term with a similar application). with their subdivisions. the subtropical. The marine biotopes mud beach. but these are so obviously exceptions that they "prove the rule. may be characterized as "fades" (originally a geological of the fauna These bear the same relation to the do subspecies and varieties to a species. climate. The depth strata of the sea are those of different degrees of pressure. climatic zones are the polar." This in turn may be united with the biochore ice desert under the superbiochore wasteland. The biotope is directly condi- tioned by the similarity of regional factors." Thus the biotope rock resert combines with sandy desert and stony desert into the biochore "desert. Certain animals enter more than one biocycle. and soil. ." appear in varying vertical or horizontal position. such as the medium. it is and the and since these are represented customary in mountains in the altitude strata to call these zones also. mountains. amphibia in fresh water and on land. and on land the altitude strata are of two sorts. typical biotope as Each of these represents special exclusive habitat conditions to the animals inhabiting it. differing from the typical conditions in recurrent minor respects. fresh water." The biosphere may be divided into three such biocycles: ocean. so may biotopes be grouped according to their resemblances into "biochores. and light (see Chapter 12). which may be called "biocycles. and land. This is reflected in the nearly complete difference in their animal populations. like salmon or eel in the ocean and in fresh water. The customary tropical. horizontally distinguished ones. Particular modifications of a biotope. sand beach. and these factors condition an analogous development and flora of a given biotope. the temperate." which with the subbiochore eroding shore makes the biochore seacoast This together with the sea bottom from deeper water makes the superbiochore benthal. we have depth and altitude strata and climatic zones. and some birds occur in all three. As species are combined into genera and these into families and orders. and they are thereby fundamentally distinct. Vertically arranged provinces are called strata or layers.gravel beach and shingle beach (boulder beach) belong together as the subbiochore "depositing shore.166 Biotopes and Biocoenoses neither can be defined with exactness. motion.. those based primarily on stratification of plants as in a forest and those based on physiographic features such as low- land. independently of other subdivisions of the biosphere.

The faunal regions and their subdivisions are based on common origin. and though they may be of joint origin are not necessarily so. corresponds with the biosphere. areas within the biochores that have uniform ex- ternal habitat conditions are biotopes. The 2 vegetation or as the phytome. is defined as includcommunities. The clearest correspondence between an area and its population is found in the biotopes. Biotic communities The sum total of living things. and land. since it takes account of ecological succession The concept and of the interrelations of land and fresh-water Thus the "spruce-moose biome. and their populations are homologous. ocean. called the biota when both plants and animals are included in plants considered with regard to their ecological relations rather than their taxonomic affinities may be spoken of as one category. fresh water. while the animal life similarly considered is called the zoome. 11 called biocoenoses. The biotopes and biochores include analogous provinces whose faunae are similar on account of the influence of similar habitat conditions. which may be united as superbiochores or subdivided into subbiochores. 1 ing the innumerable lakes of the glaciated region. vegetation type and operational overlap By continuity of it of successive communities includes peninsular extensions in the mountains of western North America and in the Appalachians even where neither spruce nor moose is represented.Biotic Communities 167 All these divisions are independent of the historically based faunal regions. cuts across Hesse's classification. subdivisions with a general similarity of habitus are recognized as The biochores. ' 17 A biocoenosis is the association of living things inhabiting a uniform division of the biosphere and correspond- . whose inhabitants constitute a well -characterized unit often called a biotic community. deserves wider use. of the "biome." extending across northern North America from Alaska to Labrador. plant and animal. and their variations are called This logical classification of habitats originated by Hesse 7 facies. Such assemblages may be split into "plant communities" and "animal communities" only by arbitrarily disregarding the essential ecological 1 Such assemblages of organisms have also been unity of the whole. in these." for the largest recognizable ecological formations. while the biocycles have their respective faunae and florae. ecological divisions of the biosphere are accordingly (to summarize) the three biocycles. on relationships among the inhabitants.

it is but The by lakes without outlets or nearest approaches to such closed systems are perhaps furnished by oases in deserts. to carry out such a division and to place consideration of the animals near the foreground.168 Biotopes and Biocoenoses ing in the selection and number of species with the average external The members of a biocoenosis are dependent habitat conditions. more or less sharply distinguished from its surroundings. the two others are not. which Biocoenoses form is self-regulating and fluctuates about a mean. but so does an anthill in the forest. 8 . for the sake of simplification. and usually food is contributed to other communities.* be considered as representing a biotope in a biogeographical sense. Since a biotic community forms a unit whose members are mutually dependent. for example of a landscape. Biogeography cannot carry subdivision to the extent that is possible for ecology in its consideration of animal communities/ is Not every biocoenosis to as a biogeographical unit presents a characteristic bit of the face of the earth. with their respective inhabitants. would be "autarchic". forms a biocoenosis. including the forest itself. the biocoenoses would form closed systems. A lower limit is set for the size of biotopes in their relation to geog- raphy. introduces aspects of ecology not particularly pertinent to the present geographical approach. But it is sometimes helpful. The dependence of the animal element on the plants is well known. without necessarily including a single species limited to an individual biocoenosis. plants are the producers. animals the consumers. it is accordingly not logical to separate the study of plant and animal elements. with interlocking interrelationships. upon each other and are thus forced into biological balance. If these mutual relations were present in every community and if the amounts of foodstuffs supplied were equivalent in both directions. The life of an oak forest. or a hazel thicket on the edge of the woods. and this limit is not necessarily valid for the biocoenosis or for the habitat niches of ecology. in contrast to a biocoenosis in which the interrelations between constituent organisms make the primary binding unity. includes an area of determinate physiognomic The biotope value. 9 * ' very rare that a biocoenosis does not require food supplies from without. characteristic ecological communities. The oak woods is a biotope. useful in a description of the earth's surface. Some biocoenoses are wholly dependent on the outer world for their food supply and may The tendency to regard a biotic community as an assemblage of species primarily controlled by the biotope.

. its plant foundation. the succession leads to chalk grazing operates. is unstable. On grassland only if the desert is encroaching upon the savannas because of grazing. though secondit determines their The protective devices of plants against browsing may exclude certain animals completely from a biocoenosis. relatively slight fluctuations about a This equilibrium. the desert is spreading because of destruction of Atriplex and Kochia by rabbits and stock. tions. mode of motion. Vegetation affords the animals not only food to the earth. but also shelter. The whole composition of a biotic community. Reciprocal effects animals are an influential part of the plant environment. so that the changes by death or increase cause only mean. of oxygen. plant metabolism supplies an abundance Carnivores are dependent on plants also. and part of the area was invaded by the cactus Opuntia. to withdraw from periodic climatic influences. It is disturbed through the variations in the habitat conditions themselves. arily. and the cooperation and competition among the animals that enter it help to determine both the species and the number of individuals in a given biotope. Within the biocoenosis. however. Animals may actually destroy the plant association upon which they In West Africa subsist. the plants are most directly dependent upon the conditions within the biotope. particularly when they are permanently present in sizeable populagrazing. 12 Similar changes can be seen in various parts of the grassland-desert ecotone in the southwestern part of the North American continent. There is a general balance. Land and shore plants are confined whose chemical constitution regulates their food supply. community in which they predominate. 16 in the arid pastoral belt of Australia between the 10-inch isohyet and desert. its competitors. the influctuations. with subsequent profound edaphic changes.Biotic Communities 169 then consist entirely of animals. The unable are They of the cover plant given biotope to a large extent determines the nature of the animal population. otherwise scrub develops. Much of the tall prairie grass in Kansas gave way to short grass under By pressure of grazing by bison. as a result of overgrazing by cattle. hinders orientation. as in caves or in the communities of the lightless depths of the sea. animals hold grasses in a preclimax stage and by overgrazing may cause a retrograde replacement of the vegetation. which undergo When the numbers of a single species change. Deer may enable the red spruce to become locally dominant by selective browsing on English chalk. and sometimes In water. the presence of the unpalatable ferns must have an important influence on the composition of a occur.

the concentration of cuckoos must have an important effect on the breeding of the smaller perching birds. flourish. Additional pertinent examples may easily be found. growth of the ground cover. as the food of predaceous anisupply favors the growth of the fry. caterpillars may leave such forests. Thus an excessive multiplication of the processionary caterpillar of the oak (Cnethocampa processioned) directly affects the oaks. the abundance of to lynxes in Canada.170 Biotopes and Biocoenoses timate nature of the internal relations within a community causes the other members to change. on the other hand. mals is dependent on the number of food animals. 13). Wild animals that take up with their food the widely disseminated poisonous and irritating hairs of the Cuckoos. such as the oak tortricid Tortrix viridana and and food are reduced. Receipts of skins of Lepus amcricanwt (above) and Lynx canadcnm (below) by the Hudson Bay Company from 1845 to 1905. for example. whose breeding places of the forest bottom by the caterpillar dung has an effect in a richer FIG. since the hairs are not objectionable to them. As the increase in the caterpillars comes in May and June. After Seton. The number depend directly on the . and the other forms dependent on them. A notable increase in the plankton of a given part of the sea has an important influence on the numbers of fishes in the following years. 13. which are eaten bare. seems abundance of snowshoe rabbits (Fig. in whose nests the parasitic European cuckoo lays its eggs. Fertilization gall wasps.

others will thrive better. Some members will drop out. The lakes of Finland and south Sweden were once part of the Littorian Sea.Biotic Communities 171 Such fluctuations in general are rapidly equalized on account of the superabundant fecundity of all members of the community. On animal account of life. in correlation with the absence of the adverse conditions found in less favored regions. members In addition to being conditioned by their biotic interrelations. climatic changes affinis. and thus have progressively more rigorous selective action. Pontoporeia Similarly. two other species that were dependent upon it disappeared with the caribou wolves and Indians. and Limnocalanus macrurus). this selective influence of the environment upon world may be cerned. and Australia are independently populated by members of each faunal area. genera. Forms that cannot survive extremes of must perish In regions (cf. has operated to destroy and drive out the native forms. species. Eurasia. American prairies. Permanent changes in the faunal composition and the numeric relations of the species result when one of the members of an association disappears or when a new one is added. in spite of the similar general impression the in- . Any permanent change in the physical environment ( as in the biotic ) conditions a change in the biocoenosis. are often regional communities. in New Zealand. 4 The introduction of numerous palaearctic species. These habitat conditions demand a higher degree of adaptation on the part of plants and animals the more they depart from the optimum. The members of the however. the pampas of the Argentine. if similar processes and conditions are consimilar biocoenoses arise Thus corresponding biotopes with formations. such as passerine birds and earthworms. optimum physical environment. in different parts of the biosphere. and still others will be enabled to enter. the communities are characterized by numerous negative features. different The most grassland areas the North entirely different. the ecologic communities in widely separated parts of the similar. Idothea entomon. The caribou has been almost exterminated in Labrador by ruthless hunting. and the entire marine fauna was reduced to a single fish (Coitus quadricornis) and a number of crustaceans ( Mysis relicta. Chapter 11). the steppes of Africa. Australia. such as tropical seas and the tropical rain-forest. since the last glacial period have completely changed the mid-European and North American biomes. analogous and the families. the of an ecological community are dependent on the physical conditions of their biotope. their separation was followed by freshening. They are. quite independent of each other. and other countries.

172 Biotopes and Biocoenoses habitants are taxonomically usually more nearly allied to those of other biocoenoses in their region than to those of the similar habitats The grassy steppe of Australia is populated by in distant regions. the South American by histricomorph rodents. in deserts. The number of these most closely adapted small for most communities. in the snowy zone of the Alps. a large of jumping types equipped with long hind limbs. heat. the North American and Eurasian by other families of rodents and by rabbits They agree in having strong chewing apparatus. These are frequently adaptable forms that flourish equally well under very different habitat conditions. the ground beetle Carabus silvestris. Tychocene animals may be of in the animal community than are the autoch- . numerous necessary differences behares. in the individual biotopes and biochores: eupolar. in the biocoenosis of moss. as ubiquitists (forms with high ecological valence. it increases with the severity of selection as the conditions depart from the optimum and is very large. One important set of animals for the characterization of a biotope are those that are limited to the corresponding biocoenoses. eudesertal. eucaval. Such members of a biocoenosis may be dis- tinguished as tychocene. Ty- when fall behind the eucene element in numbers only the selection by unfavorable conditions is especially severe. in the saline inland waters. and number tween the faunae of similar habitats result from the different degrees of adaptation to the similar conditions. in caves. in chocenous animals the tundra. in the ice-dominated tundra. and correspondingly. for example. etc. eunival. and drought are required. and a variety of burrowing forms. euvastal. and only such animals can persist as are able periodically to suspend their living functions. in the snow zone of mountains. Such animals may be distinguished as eucene. euhaline.. i. in steppes. Examples of tychocene forms are the raven.e. where resistance to index forms is long-continued cold. unable to live in another environment. whether only in similar adjacent ones or in widely scattered very different habitats. In smaller biocoenoses such autochthonous forms are fre- quently lacking. These are so exactly adapted to the ruling conditions of their habitat that they are species. since the carnivorous and omnivorous animals tend to have a wider distribution. also in other biotopes. They are usually herbivores. in the Arctic. marsupials. the index which may be characterized as autochthonous. the wolf. and all greater importance migratory birds. In addition. A much more numerous biotic element occurs eurytopic forms). awaking to renewed life when favorable conditions reap- pear.

animals are fluents. Such a climax is insensitive to its own effects and. moss. The biotic communities in the dunes along Lake Michigan evolve towards the beech and maple forest community as a climax. tramps. for the sparse fauna of hot springs consists in the main of ubiquitous forms that are sufficiently resistant to withstand the unfavorable conditions. have already been discussed. largely as a result of the activity of plants. heterotopic animals. will remain as long as the present climate persists. a few more or less accidental forms may be present. cession is effectively illustrated undergo ecological succession from Such sucin dunes where. such as the filling of lakes stantly changing. however. Such changes are accompanied by obvious changes in the Even when the climate and other physical conditions remain biota.Biotic Communities 173 thonous eucene forms. the biocoenosis may still pioneer to mature. classified as the basis of relative importance in the dominants. These are guests. and temporary pools. The biocoensis of a given biotope is accordingly the more uniform. Both the biotopes and their accompanying biocoenoses are conChanges of biotopes. other things being equal. the more severe the selection by the physical habitat conditions. This is not a uniform rule. and in general it will be the more distinct the poorer it is in species. On community. but inevitably perish if they do not find their way in time to more favorable environment. and a butterfly on a glacier are examples of such accidental occurence. influents. constant. The most distinct biocoenoses are those of deserts and ice waters. caves. with the passage of time. States. so-called climatic climax communities. and the subinfluents are regular members of the community with but relatively slight influence. or were before man's influence became too important. which live for a time in a biotope into which they have wandered. This is the climax community for much of eastern United it the gray fox and the bobcat are index mammals. or increased aridity as in the Arabian peninsula. the communities of the bare dunes evolve through various stages until the climax community of the region is reached. and subin- Dominant animals are those of outstanding abundance or conspicuous influence that are present at least throughout the entire active or open season. 15 In addition to these regular elements of a biocoenosis. a bird of the high seas blown inland. Influents are common animals of less importance. A fish in a spray pool on a rocky coast. in .

The population density must be distinguished from the species density. Coenonympha. and Pararge with 20. does not give a di eel A picture of the composition of a biocoenosis. or even mice are generally present in much larger numbers than larger ones. It must be admitted than an exact estimate of the amount of ciation is animal life in the terrestrial biocoenoses is excessively difficult to make. is well established for marine ann freshwater plankton. with good fortune he might catch a Pieris. however. from a species list. no comparable values would result. and Erebia with SO. and Epinephele are present in great numbers. which depend in turn depends directly on the biotope. greater part apply as in the associations on the sea bottom or in bird rookeries. the amount of animal life will on the amount of the plant element in the association. Erebia. the genera Satyrus with 56 species. the numbers of individuals being equal. is different in each biotope. Vanessa with 12. would be much better represented. and Coenonympha with 25.174 Biotopes and Biocoenoses Population density The amount of animal life present in an area. Epinephele. he might encounter the species semele of the genus Satyrus. or an Epinephele. midges. Pieris. It depends primarily upon the amount of food available in the biotope in question. and Satyrus species. to catch principally Sesia. He would accordingly expect on an excursion. many The knowledge insight into the of this factor. a Coenonympha. The reality stands in complete contrast. If the assoa closed one (autarchic). This factor. Such a relation does not the all of when or food the comes from without. Pieris. in an area of considerable size. and associations between larger and smaller animals should be richer than those between small forms alone. An illustration written in 1890 makes the point: "One would find Pyrameis with 4 species. but it would be too much to expect that he would meet with a Vanessa or Pyrameis. The former is measured directly by the mass of animal substance present. Sesia with 78. since small animals such as hydroid polyps. butterfly life of the palaearctic fauna gained from it species catalogue would be quite wrong. The amount of animal life in a given biotope may be considered from different viewpoints. whether they are abundant or scarce. which may be termed the bio-mass. Species density. the . on the other hand. but of u the 78 species of Sesia he could not expect to find one!" Whereas the population density represents an absolute value. is well known in This is the number of species that arc encountered regions. If it were to be estimated from the numbers of individuals.

The ." says Mobius. The utilizato tion of vegetable food requires certain adaptations and is not possible for all animals. insects. All of the usable food-materials present in it will be claimed by the life produced 11 This applies. especially islands. 10 Only a very few adaptfavorable. the animal population density.Population Density species density is a relative one. In equal areas with similar conditions of food supply the total mass i. able animals inhabit the ooze banks of the shallow North Sea. which are necessarily euryhaline on account of the varying salt content of the water. herbivores are only sparingly represented. be less the species density is very great but decreases as the environment be- comes less favorable. however. The principal herbivorous groups of the land are the land snails. Species density depends on one upon the selective action of the environment but on the other historic relations.175 hand upon Thus the species density upon an oceanic island than that upon an island of equal size near a continent. such as the remaining groups The coloniza- New Zealand. though in small variety of species. and mammals. with a greater wealth of species. birds. and others. tion of several regions. but. Copepoda. the Falklands. of most a number of have smaller the polar seas species groups of animals than the tropical waters. the number any one is smaller. other conditions being equal. Two biotypes with equal population density may be very unequal in species density. will in general . Plankton catches are mostly quantitatively larger in summer in the arctic seas than in the tropics. to the extent to which the members of the biocoenosis are able only make use of these organizable substances. conditioned by historic factors. of life produced. with hoofed mammals has shown how much more animal life they were able to support than was actually present on them. From this results the peculiar inverse proportion between Thus of species and number of individuals in the species. This is a rather direct corollary oligochaetes of the seacoast. are present in very great numbers when food conditions are of Mobius' proposition. but single species of Radiolaria. "Each biocoenotic area has the greatest mass of life in every generation that it is capable of producing and supporting. there. but not a smaller population number density. The full use of the food materials prepared by the higher plants of a region may be prevented by the absence of animals. animals able to utilize the available food supplies were absent. of individuals of while in the tropics. especially for animals. Under optimum conditions. In all of animals. and fishes appear there in enormous numbers of individuals.e. will not be very different..

and often. which may be short. a four-year period in north Germany. The factors." Zool Anz. "Grundsiitze und Grundbegriffe der biocncnoiischcn Forschung. Press: viii. 8 In favorable regions development It is quite otherwise in the tropics. Franz. Generation follows generation. those that die only follows. necessitated by the winter.176 Biotopes and Biocoenoses few animals are present in enormous numbers. The principles of animal ecology. resting stages. (vol. 2. 1910. 306. Orlando Park. C. but whether the fauna will be uniform or varied depends upon other The Newly developed forms are more likely to be preserved under optimum conditions than where the selection is more severe. Here life pulsates with much more rapid beats. The European may beetle Mclolontha vulgaris has a three-year generation south of the Main. Oklahoma Univ. xii. and even if death comes earlier." Begriffserklarung Viertellahresxchr. R. 837. naturforsch. they may be present in such numbers as to color to a deep red broad areas in shallow water. Das Tier als Glied des Naturganzen. 63:293-493. and in east Prussia its development takes five years. and in birds and mammals as well as in other vertebrates the breeding period is often not restricted to any season of the ) ear. Gams. which feed on the rich deposits of food materials.. available food accordingly determines the population density. 4. Teubwr: xv. 5. 1949. Leipzig. 3 maps. H. the high temperatures hasten the development of the poikilothermal forms. l\ Schmidt. 33:349-353. Tierbau und Tierleben). 1918. Friedrich. Doflein. . Zurich. and Karl Philadelphia. "Principienfragen der Vegetationsforscbung ein Beit rag zur und Methodik der Biocoenologie. Alfred E. the forms. make room for the new generation that BIBLIOGRAPHY 1. An ecological glossary. The development of organisms is slow. Norman. as in insects.. Alice. 960. J. Thomas Park. 263 figs. 1938. In the temperate and colder regions active life is concentrated into a certain season.. 2 of Hesse and Doflein. Emerson.. Variety in the plant world also favors variety of animal life. as the unfavorable conditions of this habitat prevent pursuit.. Dahl. Ges. W. 20 pis. Saundrrs: Carpenter." The slimeworms Tubifex. 3. may extend over several years. interrupt the active life of the great majority of animals and probably increase the longevity of the individual. as in an insect whose life is closed with the conclusion of egg laying. 1908. is scarcely or not at all interrupted. share this biotope with only a few other as these banks are rich in food matter. 740 text figs.

"Animal communities of an (p. Ecology. 135 figs. 17. 288 ff. 1937. Teubner: ii. 3:665-682..f. 8. 10 11. Die geographische Verbreitung der Oligochaeten. J. Budapest.. ("The oyster and oystereulture." Illinois Zool prairie. Bibliographisches Institut: 2 vols." Naturw. Richard." Rept. map figs.. Friedrich. 5:281-343. 1937. illus. sylvic. 298). (n. F.Bibliography 6. Leipzig. Shacklcford. & Parey: 126.. Fish Comm. 1880:683-751). "Allgemeine 1 Biologic der Schmetterlinge. Ernst. 70:1-28. Hesse. 1. illus.. 1890. sectenkunde 9. P. illus. 1914. "The problem of the Sahara in West Africa. Australian Council Sci.. Berlin. 12. N. with special reference to southwest Queensland. ..). Holzcl: 2 vols. maps. Stubbing. W. Ehrenbaum.. Syst. Vienna. Wocfischr. 7. Judeich. 10. 1 map. Ratzel. W. kunde. Heuipcl. (vol. Research Pamphlet. M. 13. S. Tiergeographie auf Oekologischer Grundlage.) 17:281-304.. and H. 186. Kjellen. Jena. Rudolf. Die Awfter und die Austernwirtschaft. 10:126-156. Seitz. 613." Ind. Die Grossmdchte der Erde. Michaelson.. Friedliinder: vi.. 1903. Adalbert. 208 (p. 1901. 1895. Lehrbuch der mitteleuropdischen Forstineinem Anhange: Die Forstschddlichen Wirbelthiere). 1 pi. Mcibius. Nitsche. Wiegand. 1877. 14. Karl. Thienemann. p.." Acte* ifvme congr. 1924. 10 16." Jahrb. (mit Berlin. Seemuscheln als Nahrungsmittel 1924]. 9). "Further observations on soil erosion and sand drift. 177 [fide Hesse. intern. 3rd ed. E. RatclifFe. 1918. maps. "Lebensgemeinschaft und Lebensraum. Die Erde und das Leben Eine vergleichende ErdLeipzig. F. 15. U.. Fischer: xii. 1929. 9 figs. August.

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however.000 by land. present about 3. would be everywhere of the same depth. or 361. more than two-thirds.000.. are occupied by the oceans and only 149. * THE OCEAN PRESENTS THE MOST EXTENSIVE HABITAT FOR LIVING ORGANIf the total surface of the earth is reckoned at about 510. and Fleming (1942). It falls into two main divisions.000 cu.000. whose title. If the level of the sea bottom were equalized so that the oceans.000 sq.000. T/ii.000. In many ways it is convenient to discuss hydrobiology as a unit instead of dividing it into its oceanographic and limnological aspects.370.000 metric tons of material from the land are being washed into the sea annually.000. are more easily dealt with separately. yet Welch that only about 2% of the land surface of this continent is estimates covered by inland waters (see Chapter 16). If all the land were submerged in the At sea. the Atlantic with the Arctic on one hand. Such waters occupy only a small fraction of the land surface. km. only a relatively small amount of water would be displaced. The oceanic mass of water forms a single continuous domain.) 179 but conceals . suggests its readability 1* its accuracy. present a more detailed summary.' (Citations are given at end of Chapter 10. including adjacent seas. lakes and has :>T at least * The his general problems of oceanography are discussed cogently by Coker in 1947 book. In contrast the mean elevation of the land is only about 700 m.000. From the zoogeographic point of view.. from which the volume of water contained in the ocean basins may be reckoned at 1.Part 2 The Distribution of Marine Animals INTRODUCTION isins. whose physical and biotic characteristics constitute the subject matter of limnology. North America is rich in an average river and stream surface. inland waters.v Great and Wide Sea. in The 84 Oceans. Johnson. Sverdrup. that depth would be 3795 m.000. km.

in thickness. species something like one-fifth are aquatic animals. whose inhabitants are constantly. all parts of the ocean.. does not ordinarily exceed some 30 m. In the ocean. Its tinuously. equally larger number of inhabitants. This vast space is everywhere inhabited by living organisms. or The atfor the greater part of their life. such as the Mediterranean or the Baltic. they are connected with the principal oceans by straits. The separation of these incomplete. A division like that of the land into continents like the Aral is thus entirely wanting for the oceans. which. Terrestrial life occupies but a thin stratum. and their waters are united by wide stretches at the south. but living animals have been brought to the surface from even the greatest depths. and Caspian. When this is contrasted with the mean depth of the oceans of almost 4000 m. of species is less From this it may be concluded This has its that the evolution favored in the sea. and may be represented in all their stages at all depths.180 and the is Marine Animals Pacific and Indian oceans on the other. . even in forests. and at the north by the narrow opening of Bering Strait. confined to the ground.000 air-breathers. contributes so much to its evolution into new made difficult by the confluence of is favored on land. mosphere is not a true habitat factors in the life of land animals. Very few animals are found in the deeper regions. much reason in the great smaller development is The isolation of a group of animals. it is occupies more than twice the surface area seen that there is more than 300 times as much readily space available for marine as for terrestrial organisms. whereas such isolation Despite the smaller number of species in so much greater a space. for living organisms or their rather. and when we remember that the ocean of the land. which forms. It contrasts with the atmosphere. one of the ecological low density makes it impossible developmental stages to float in it conit is. While a number of smaller divisions are essentially independent seas. are few and vanishingly small compared with the oceanic area. and air-breathing animals are therefore superficially distributed and in principle confined to the earth's surface. It is true that the strata are not in dense the upper ones have in general the population. In spite of the much greater extent of their domain. out of 822. including the fresh-water forms. Completely separated salt-water basins. living organisms are permanently suspended. uniformity of habitat conditions and the of barriers to distribution. the number of species of the inhabitants of the sea is much smaller than that of the On the basis of Pratt's figures of 1935.

33 the is much Onychophora. and Amphibia. the Mollusca. the Amphibia. Marine representatives of the inverte- brate phyla are already present in the earliest fossil-bearing strata of the Palaeozoic. Chapter 4). Among the vertebrates. On the other hand. Diplopoda. are wholly wanting in the sea. in the Devonian. still This set of relationships is most simply explained by the assumption that the ocean is the original home of life. 25 classes. and a weighty argument in its favor lies in the fact that the body fluids of the marine animals (with the sole exception of the highest and most recent forms.Introduction the animal life 181 that of land and of the sea exhibits a greater diversity of form than does fresh water. as compared with the 4 air-breathing classes. in the late Cambrian. by way of exception. only 4 classes out of the total of 59 recognized by the 1943 text of Storer. on account of the different conditions of fossilization on land and sea. require special modifications to prevent the dilution of their body fluids by the water in which they live. The explanation of this fundamental diversification of all marine life lies in its much greater age. and the predominantly marine Cyclosto2a referred to under the genmata. great concentration of their body fluids in consequence of the loss of water (cf. so that no osmotic extheir body fluids and the surrounding Fresh-water animals. . and the first air-breathing vertebrates. and terrestrial forms require protection against the too change takes place between medium. Enteropneusta. the bony fishes) are isotonic with the sea water. It appears certain that this relation in the marine forms is the original one. on the contrary. and that in the other two we are dealing with new acquirements or "adaptations. When a phylum has representatives in fresh water and on land as well as in the sea. including the entire phylum Echinodermata. are purely marine. or the Chordata with the exclusively marine subphyla Acrania. and several other classes eral heading of fishes. Selachia. Disregarding parasitic types. Chilopoda. the air-breathing classes are more numerous than the marine." The waters of the ocean approach the ideal medium for living substance. although it is possible that the remains of terrestrial animals of the same age have not been found or have not been preserved. with only the Pulmonata and a few Prosobranchiata on land as compared with the varied marine classes. Among the Arthropoda. the fishes appear first. the former tend to be less varied in form than their marine relatives: thus. The air-breathing forms appeared later and one by one. and Tunicata. thus in the Palaeozoic. the variety of structure among marine animals greater. for example.

. modern out plated direct counting methods show that the bacteria are from 200 to 1000 times as abundant as was indicated by plating/ 16 Waksman has found that in the Gulf of Maine in water from 200 to 350 m. among others.000 to 347. Both and the anaerobic Azotobacter Woods Hole are nitrogen-fixing organisms. and they have been reported as being entirely absent in some of the samples. at least by the methods used. ordinary uncontaminated sea water in nature appears to be a relatively poor medium for bacterial growth.) re- which has been com- bottom deposits and near land. Earlier investigators their samples in order to estimate numbers present. 153 of ref.000. by Waksman and associates using the facilities of the Atlantis of that abundant populations of the aerobic Closteridium occur in the sea. least important. but they are more abundant in the water immediately above sandy as compared with muddy bottoms. In fact.000. when contaminated areas are avoided. Still fewer bacteria are contained in bottom materials from oceanic depths." Waksman sug- monly found in gests the following hypothesis concerning the nitrogen cycle in the 86 sea: .182 Marine Animals role of bacteria in the general The economy of the ocean is still a matter requiring further investigation. the sea water itself. Off Andros Island as many 8 as 160. There are three centers of bacterial in the sea: the marine plankton. The number of bacteria on the continental shelf decreases with distance from land except in the deeper layers of mud. where they remain constant.000 bacteria were found per cubic centimeter of mud. The long dispute concerning the presence or absence of nitrifying bacteria in the sea continues despite conclusive evidence produced. and. there are other marine nitrifying organisms which have escaped detection. of water.000. Apart from their food value. Bacteria are present in fewer numbers on or in sand.000. of which we know practically nothing for the ocean in general. bacteria in the sea are chiefly concerned with the decomposition of biotic of certain simple life residues and with the transformation compounds or elements. They live both at the upper surface and in decreasing numbers deeper in the mud. the sea bottom. The differences on the depend type of plankton rather than on depth or other factors. and on George's Bank in 60-75 m. ZoBell in 1946 cords his belief that "besides the soil-like nitrifier 39 (p. the numbers of bacteria per cubic centimeter of plankton tow ranged from 202. in the sea 35 the ratio between numbers in the tow and numbers water alone was from 225:1 to 2270:1.

On reaching the zone of photosynthetic activities. Reduction of nitrate to nitrite does not mean necessarily any loss of nitrogen from the cycle of life in the sea. . Very is nitrate reduction to gaseous nitrogen or complete denitrification is possible under normal sea conditions. where nitrate it remains as such. This ammonia is rapidly oxidized by specific bacteria living in the bottom to nitrite and later to nitrate.Introduction 183 Decomposition of the organic nitrogenous compounds takes place in the sea water but largely on the sea bottom. due to a lack of available energy for the nitrate-reducing bacteria to a lack of such bacteria. This nitrate remains in the sea bottom and is and not not reduced. which may also be gradually consumed by the plants. for the greater richness of life in the shallower waters of the continental shelf. The small amounts of ammonia found in the sea water originate from the plant and animal residues in the plankton and in The nitrate formed in the bottom gradually diffuses into the water the water. in addition to later. little By such those to relations there are seen to be good reasons. and in polar regions be given where the circulation tends to bring the nitrites and nitrates up to the lighted zone. with the result that the ammonia is then liberated. on oceanic banks. this consumed by the phytoplankton or is reduced by the nitrate-reducing bacteria to nitrite.

.

which may afford a means of characterization of certain communities. that falls The amount is of reflected light back reaches a minimum of 3 to 4% in Buzzards Bay near noon on clear calm days. Apparently. similar changes not infrequently appear in animals of similar habitats. Physical in Conditions to in the Ocean Relation Animal Life MARINE ANIMALS ARE INFLUENCED IN MANY WAYS. It increases a few units when there are waves and a few more on cloudy days. amount reflected of the rays. IN BOTH STRUCTURE and activity. partly by adaptation to them by means of selection. Partly under the direct phenotypic effects of these influences. As the sea's surface becomes more disturbed. is greater with greater slant that the the fact this must be added absorption occurs from the surface Slanting rays do not reach as great a 185 . The surface loss is usually about 15%. the "surface loss" of light becomes greater although the reflection is increased but little. To in the direction of incidence. by the physical and chemical properties of the surrounding medium. These influences of the medium concern and us here only in so far as they inlife. treatment will be more or less detailed accordingly. under these conditions the amount of light absorbed in the upper meter or so of depth becomes larger. It has also a primary influence on the coloration of ani- mals and upon their organs of vision.10. fluence the evolution distribution of animal The importance and their of single characteristics in this respect is very unequal. although it has often been reported as reflected on the surface of the sea and being much 5' 30 The greater. Light The amount of light in the different strata of sea water determines primarily the development of plant life and thus influences animal life secondarily.

at a given time. i. and 25 m. In fact. and thus varies with location. the green more abundantly. as at river mouths. at 610 m. while even 40 minutes' exposure of suitably prepared plates fails to discover a trace of red or green.e. Beebe. the photographic plates are unchanged diving in a bathysphere near the human eye at 579 m.. but the red rays are most sparingly represented. little light with wave lengths less than 6000 Angstrom units (A. 25 Traces of light can be detected at a depth of 1000 m. rapidly than the short. the water is quite different from that in air. The composition of light in all evidence of sunlight had vanished. at 67. Yellow or orange rays penetrate such water most readily. The different wave lengths composing ordinary light are differentially absorbed by the water. At 500 m. in the open ocean in the subtropical zone by means of photographic plates. The depth at which light is reduced to \% of its surface value is found at 8 m. The amount of light at this depth is one three-millionth of the amount at a depth of 1 m.. Beebe in his bathysphere dives detected no violet with his eye. Animals at intermediate depths are frequently red in color and presumably appear black in the absence of red rays. is about equal to that at 500 m. but only a strange blue that appeared brilliant even after losing power to illuminate objects. after two hours' exposure.e. at 33 the same position of the sun (i. at 50. at the same moment of time) the light at 800-m. . and only about a quarter of an hour at 40 m. all the colors are still present. while blue and violet predominate. 2 Bermudas. At 1700 m..186 Physical Conditions in the Ocean strike depth as those that more nearly perpendicularly. 40 m. N. With is about 50 m. 1889) had a length of 11 hours at a depth of 20 m. The vision depth. and. Toward the poles the light falls at a lower angle with the surface of the water and thus in general reaches a lesser depth.. Monaco observed that the "day" in the water at the harbor of Funchal (March. Thus the amount of light reaching a given depth at a given place varies with the time of day and year. the depth at which a Secchi disk disappears. found light visible to the the greater is the amount of this difference. and the deeper the water. Transparency of sea water also depends on the amount of material in suspension. at 67. orange) penetrates more than a meter into strongly colored water. differs according to the latitude of the place. depth at 33 N. in the Woods Hole Harbor.. In clear water the long wave lengths are absorbed much more At 100 m. at 50 and to that at 200 m. 5 hours at 30 m. the blue to ultra-violet rays are still abundant. latitude. whether organic or inorganic. according to the presence of large amounts of plankton or of finely divided silt.

according to Lo Bianco. Murray and Above 200 m. reaching depths of 4-5 m.. rich in light. in a still is notable in coastal waters. Despite some apparently as the result of the attack of a protozoan parasite. Transparency decreases greatly on the continental opacity of fresh waters is shelf. where light is excluded by 40 to 50 m. reached depths of as much as 40 m. at 75 m. eelgrass had been greatly reduced on both sides of the Atlantic. it was even more limited. according Murray and Hjort. to 2. with the possible exception of the Mediterranean. 1*' 5 The differential absorption of light exerts an important effect on the distribution of plant life and its dependent animal life. of the Sargasso Sea. where the light extends to a lesser depth. according to Lo Bianco. Even before being much reduced by mycetozoan * attack. The same effect. in most in the fiords of Jutland. and these are present in sufficient amounts only in the uppermost levels. The reduction of the amount different depths. 80-200 m. down to 500 m. is better realized when it The relative known that the clearest yet measured. according to to 100 from Chun. in the transparent waters The transparency of the Sargasso Sea is the greatest yet measured. and the maximum is approached at the depth of 10 m. signs of recovery there is no certainty whether eelgrass reduction is temporary or permanent. The terms introduced by higher degree.Light at 187 32 m.. The euphotic in stratum. in the North Atlantic. The waters of the ocean have been divided into three strata with reference to the amounts of light present. Schimper. * . Wisconsin. extending from from to 30 m. orange. The dysphotic stratum. this corresponds to the "shadow flora" of Hjort. It is precisely the red. is approximately equal in transparency to the water of Vineyard Sound. and at 149 m. only one-fifth. which in turn ap- proaches that of Woods Hole Harbor. By 1949. Plant distribution is accordingly very unequal at of plant plankton comes between depths of 20 and 50 m. rich in whereas coastal waters.. in the Gulf of Maine. 30-500 m. the Danish coast. Chun are: 1. 80 in. rich in phytoplankton to 1J) and herbivorous animals. and at 100 m.. transparent It plankton. Crystal Lake. and yellow rays that are most important in photosynthesis. the eelgrass Zostera grew only to a depth of 14 m. there is only one-half the amount at 50 m.. weakly lighted. according to Chun. m. on below Seaweeds and extend algae scarcely turbidity. The maximum development of plant life with depth occurs still more rapidly in polar waters.

of average density. 9750 in. is not an invariable condition for marine life. are so abundant in both individuals and where their food is both accessible and con- Density statics density of the oceanic waters has an important influence on the* The density of living protoplasm is of marine organisms.. an assumption be confirmed by the investigations of Edward Forbes Aegean Sea in 1843. in height. below 1700 m. dependent on The more marine life for their species in the polar seas. Finds of living animals. Physical Conditions in the Ocean The aphotic stratum. tection. however. Thus the strengthening of the body for support and pro(Chapter 13). so that it sinks slowly to the slightly greater marine bottom.07 m. In one of the greatest oceanic depths yet known. that such an enormous pressure must crush all living beings and that the greater depths of the ocean must be lifeless. below m. nearly the poles are approached. Murray and Hjort. the pressure at the bottom is 962 It was formerly believed atmospheres.. of than sea that water. As the special adaptations for motion in open water lead to various convergent transformations. or 731 meters of mercury. however. which characterize pelagic life. This perhaps explains why the sea birds. so that only minor supporting structures are required. according to Chun. food. and the removal of this limitation helps make possible the variety of structure in the sea. north of the Tonga Islands. gradually became more frequent. perhaps on acthat seemed to in the . exerts a pressure of one atmosphere. and departure from the pressure in the sea roaches an enorat sea level is much greater than can occur atmosphere. which is required by terrestrial animals. A column of sea water 10. the shallower is the and the closer to the surface is the main mass of stratum euphotic marine life. animals. from more than 2000 m. and though none have yet been taken in the greatest known depths. 500 according to Lo Bianco. it is lightless and without herbivores. these will be examined in more detail in the discussion of that fauna The density of the sea water acts essentially as a to the animal support body. centrated. are enabled by various Many The arrangements to equalize this difference and to swim or float in the water.188 3. only detritus-eaters and predaceous forms are present. below 200 m. Pressure The pressure of water in great depths of the mous figure.

any rate The many A considerable number of pelecypods and some snails range from the surface to 2000 and even 4000 m. Any impoverishment of animal life at such depths results from the scarcity of nourishment and not from the pressure. the presence of life at depths between 6000 and 7000 m. Steel cylinders filled with water from the greater depths did not burst by the expansion of the contained gas with the reduction of the outward pressure. in the bony fishes The assumption without an opening to the swim bladder. species of animals that have a great vertical range in the sea may be referred to as eurybathic. in dredging from 1650 m. At any rate. has been fully confirmed. These animals live under a pressure of 600 atmospheres. down to the to tion. as was expected. It is true that. Thus. 9 . is not borne out.or 40-fold variation which contrast with the air-breathing animals. The rapid reduction of pressure seems have less to do with this than the difference in temperature between the depths and the surface. level they habitually maintain. 12 are to be found also within the 200-m. extreme that greater amounts of gas are dissolved in the water at great depths.. and at from the With the exception it does not prevent the existence of life at great depths. or at least greatly injured. to return at daybreak to the many and pelagic fishes live animals of the open sea. where in contrast with the ocean a uniform temperature of 12. Among 20 annelids that reach depths greater than 1800 m. and more rise vertically at night. They are not in affected by a 30. for quite a reduction of the atmospheric pressure by one-half produces injury. by day at greater depths. but as the same pressures exist in their count of the difficulty of body fluids. when they reach the surface. line. and that this creates changed conditions for animal life. often to the surface.Pressure dredging operations.9 rules from a depth of about 160 m. the pressure factor apparently does not play an important role in the life of marine animals. of pressure. the animals of the depths reach the surface in good condi27 Great variations of pressure in short periods are the daily experience of Many plankton animals depths of 400 m. In deep-sea dredging it is the general experience that almost all the animals from great depths are dead. the gas in the bladder ex- pands to a great degree when the fish is brought up from the depths. among the former. in the Mediterranean. and may project from the fish's mouth or even explode and burst Such fishes are probably capable only of gradual changes its body. in correlation with the increased pressure. bottom. of these fishes. there is no possibility of their being crushed.

and the reef corals (Madreporaria). irregular in outline in (Patella) 29 exposed places than in sheltered ones. or they may anchor themselves by means of glandular secretions like the byssus are in part direct and in part secondary. and currents and nature of the marine faunae are influenced in The distribution the highest degree by the varied movements of the sea water. Animals exposed to moving water on rocky coasts are thus in danger of being crushed by the water or torn from The inhabitants of rocky their places and hurled against the rocks. tides. with its variations salt content and its influence on the suspension of organisms in the water. Purpura.5 in one of the same size from the protected Bay of Kiel. Waves. 7 Examples of the contrastingly stenobathic forms of shallow waters are the mollusks Patella. like the barnacles. and Mytilus. the annelid Arenicola. or 1.5 kg. caused primarily by constant winds. life The shells of the limpet of certain mussels (Mytilus. while the same species in the quieter . among snails. Haliotis). per sq. then there is the struggle for existence and perhaps other causes as yet unknown.190 Physical Conditions in the Ocean is found from to 3250 m. The effects of these movements of the water upon its animal The Engineers reckon the average force for the North Sea at 15. will be discussed in connection with the pelagic fauna (Chapter 13). and the selachian Chimaera. which result from cosmic causes.4 Many species of sea urchins on exposed coasts bore holes into the rocks. smaller. for example).000 kg. tides.. and Scrobicularia longifrom 35 to 4400 m. thicker. Temperature Modiolaria discors callus probably of greatest importance. Haliotis. more and are lower. alike by the waves that break as surf on the beach. augmented by differences in density in different latitudes. m. They may be grown fast to the rocks. Natica groenlandica ranges from 35 to 2350 m. as compared with 26. coasts within the surf line must therefore have strong protective covering and must be able to attach themselves tightly in at least one of a number of ways.. the mussel Limopsis. is The from temperature changes and internal friction or viscosity of sea water. Stenobathic forms of deep water are represented by the snail Turns. and the more or less constant oceanic currents... pounding force of the surf may reach high values. A Mytilus shell from the wave-beaten west coast of England may weigh 58 gm. It is likely that pressure relations have little to do with this limitation. per sq. cm. or hold themselves by means of a powerful suction apparatus like the foot of the chitons and many snails (Patella.

20 Corals in moving water tend toward a rounded or flattened and thick mass. it is much impoverished after continued The north winds. which drive the surface water out of the bay. Tides. for example) does not make use of this means of protection. especially in the crests of waves and in surf and spray. with relatively small local variations. and in summer they carry warm-water species like the siphonophore Physophora hydrostatica as far north as the Lofoten The free-swimming stages in the life history of many fixed Islands. the larvae being brought Bermudas by the Gulf Stream. corals.Waves. on the contrary. solution of from the the increase of the favors motion air. The hydroid polyp Bougainvillia in is ramosa. in lesser depths where the wave action becomes effective at low tide. Wave motion in the open sea extends to depths . ing water the stalks become stouter and were thought to represent transformed into ramosa. with the single polyps In quiet waters the stalks are closely attached to the upper side. 12 The delicate Alcyonaria lightly calcified and finely branched. they produce the general mixing of the water and maintain the average chemical composition of the sea water In the first Wave a uniform level. but extend to considerable depths. Continuous currents contrast with the oscillating motion of the waves and tides and are of the greatest importance for the distribuCurrents carry characteristic forms of the tion of marine animals. slender. or slow-moving animals. such as the appendiculates Oikopleura labradoriensis and Frittilaria borealis. like hydroids. they have stiff skeletons. and echinoderms. and Currents 191 seas (Strongylocentrotus in the Mediterranean. may also carry animals away in definite directions. Thus in the Bay of Naples. fruticosa was experimentally Similar transformations have been demon3 Campanularia Integra and Sertularella tricuspidata. where the surface fauna after to the the sirocco is especially rich."" Thus the gorgonid-faima of the Bermudas is almost completely West Indian. The motions of the water are not merely superficial. indirectly of extreme importance for the place. where the water 16 is quiet. is blown into enormous masses in sometimes atirita Aurclia jellyfish the river mouths of the south shores of the Baltic by the summer marine winds. 10 strated in a distinct species until the supposed B. delicate. and branching. in greater depths. by oxygen at surface area. cold-water plankton. Ocean currents are fauna. caused by continued winds. (Spongodes). are elastic and wave back and forth like grain fields in the wind. are carried great distances by currents. 17 Occasional surface currents. In movdeeper water. from the Arctic Ocean into the North Sea in the spring.

A compensatory movement of the deep water must equalize the The effect of the surface currents. especially of the trades. some places. rents. both up and down. 23 The socalled "tongue of cold" in the south equatorial current north of Ascension Island is supposed to be produced by the rise of sea water . continued west winds blow the surface water away. they will be discussed at greater length in a subsequent chapter. not below 40 m. where it accumulates. a cable near the Canary Islands in depths posited of 1800 to 2000 m.5 miles per day. with their colder water from the depths. This coincides with the usual lower limit of the continental tidal currents in many places appear to extend to great in 10 years at least 25 mm. in parts of the Bay of Naples a heavy dredge may be 18 The curlifted and supported for some distance by such currents. at a depth of 1800 m. and the rise of the water from the depths is noticeable on account of the higher salt content. especially in the tropical east coasts of the continents. and the whirlpools of the Straits of Mesin sina are famous. the "mud line. It is replaced by the upward flow of cold water on the lee coasts. which is open to the east. 43 species of deep-sea fishes have been collected on the beach at the lighthouse. per month. 2 and other deep-sea forms are found at the surface there. are of especial importance in the mixing of the sea water. was found to be entirely free. On the lee coasts within the influence is continually blown away and driven against the windward coasts. These currents appear to be caused by the differ- ence in tidal phases in the Ionian and Tyrrhenian seas. and are less abundant on the surface at 15 the west coasts. on the Algerian coast.. The rate of upwelling near southern California is apparently about 20 m. which is characteristic of the deeper water of the colder and hence denser waters of the Chapter 15).192 Physical Conditions in the Ocean of several hundred meters. polar seas sink and flow slowly along the ocean bottom toward the The rate of flow at great depths has been estimated at equator. The warmth-limited reef corals accordingly thrive. The same was true of rocks on the sea bottom off the south coast of Ireland. the warm surface water in bays. Baltic (cf. and 200 m. They have been observed with certainty The vertical currents. in the open ocean." is usually given as marking the extreme lower extent of wave action near land. it reaches a lesser depth near the coast. shelf. which reaches or even 14. The about 1. of globigerina ooze is dewhereas depths. so that counter currents meet in the Straits of Messina. both up and down. In the Bay of Kiel.

This is one of the principal means of heat transport. Descending currents seem to be present in the Sargasso Sea and perhaps in similar centers of oceanic drift. Regional temperature difof are ferences importance as barriers and as impetus for variation (Fig. 14). ture reaches 34 in summer. layers. of the oceans is much to regulate the temperanine-tenths of the heat surplus Approximately used in evaporation and is stored or transported in until released by precipitation. Heat and temperature The oceans are a great storehouse of the heat that falling comes ultimately from the sun as a part of the radiation on the earth. and on one-third of this (especially in the tropics) less than 2. whose nature and origin are as yet obscure. In addition. whereas seasonal variation is high in the temperate zones. in the Baltic 17. Variation is also high in arms of the sea in the The annual temperature range in the northern higher latitudes. soma heat from the land is carried to the oceans.Heat and Temperature 193 from the bottom. The amplitude of seasonal variation in temperature depends on the location. perhaps after considerable delay. particularly to their bays and gulfs by the inflowing waters of rivers and smaller streams. in different degrees. The annual variation in temperature is than 5 on almost three-quarters of the oceanic surface. in the inner parts of the Yellow Sea 27. far back over the land. The rate and direction of all these currents are subject to continual fluctuation. The temperature of the sea water varies not only with location but also water vapor with the seasons. although even in the tropics the surface water of about 34 in the Red Sea the temperathe open ocean does not exceed 30. 2 * The vast oceanic reservoir of heat does ture of the world. A stratification of it whereby a result of the property of water increases in density as the temperature drops. Mediterranean reaches 14. in part with a kind of periodic pulsation. since they occupy much more than half the earth's surface. The combination of all these motions 25 produces a phenomenon of extraordinary complexity. ceasing at Such seasonal variations apply only to the surface depths of about 200 m. The heavy temperatures is . Temperatures of the tropical and polar seas are relatively uniform. The majority of this insolation strikes directly on the seas. The extremes range from 3 to 42. perhaps over the sea or. The greatest variations appear in localities where warm and cold currents meet and less predominate by turns. Temperature relations of the water are of maximum importance to marine zoogeography.

for 1.44 is 1. the freezing point is depressed.28 1. salinity *Per when discussing . it is approximately imum temperature at the mille (%*) has obvious advantages over per cent (%) of the ocean. Degrees Degrees Meters 183 ' Centigrade 15. sea water with a salinity of 24. temtemperatures of peratures of the abyssal waters are usually slightly above zero. 14.9. and the effective distribution of heat in the depths accomC C Winter Spring Summer Autumn FIG. according to Murray and Hjort.05 2012 2.95 Meters 1097 Centigrade 3.89 366 10.7 per mille (% f.78 Over 80% or of the ocean floor a mile or more helow the surface and has a tempera- ture of 3 less. Temperawater with a salinity of 35%o. In the sea. bottom 1 may occur in regions with polar currents. and the lighter warmer water of the tropics flows toward the poles.83 549 732 * 2743 4023 5. of Scotland Variation of the surface temperature in the open ocean off the west coast After Meyer and and in the stagnant Baltic Sea ( ).05 7. plished by currents. The radiant heat of the sun does not reach deep into the is water. On account of the presence of salt. 38 Unlike fresh water. a property limiting the min- * bottoms of lakes to 4. ) ( Mobius. The average temperatures at different depths for 25 are: the oceans as a whole.194 Physical Conditions in the Ocean cold polar water sinks and gradually spreads out on the bottom toward the equator. Fresh water becomes lighter below 4.) or more continues to become heavier until its freezing point is reached.

After Murray and Hjort. to the deep22 In est part of the sea (3968 m. such as the Red Sea.000 miles apart. with the isotherms of the sea water. points on the bottom 10.17. Diagrammatic vertical section from west to east in the sea at the Straits of Gibraltar. 15. on account of more rapid evaporation and smaller influx of fresh waters. Special conditions are found in seas connected with the adjacent tance of si* miles or ocean by more or less shallow straits. the Sulu Sea the temperature of the water is 10. the Caribbean Sea. has a depth of 1300 m.9. the Gulf of Mexico. their greater density causes a sinking of the surface waters. in Sea and the Gulf of Mexico to have a m. the Caribbean 1100-1200 ' uniform temperature of 4. Here the strait between Cuba and Santo Domingo. which keeps out the colder waters of the Atlantic. The deeper sist of the broader superficial inflow and a deeper outflow. of temperature in the sea is not uniform with increase in the depth. and the Mediterranean. As the Mediterranean waters have a higher salt FIG.Heat and Temperature ture differences 195 between bottom and surface are at the equator. found the water in the depths below A.) is uniform at 12. corresponding with that of the lowest level of the inflow. Agassiz to the bottom. the Sulu Sea (west of the Philippines). waters of the Mediterranean accordingly have a temperature of about 12. colder water does not enter. 15 ).89 (Fig. depths in meters at the left. content than those of the Atlantic. The Straits of Gibraltar have a greatest depth of about 400 m. a dis- much greater than those between two less. the temperature from about 160 m. which are replaced by an inflow of the The countercurrents in the straits conless dense Atlantic water.5 from about 750 m. At a certain depth the drop in temperatures is The gradation .

of course. The response of marine animals to temperature is naturally varied. the present temperature zones appear to be a relatively unusual world condition. which endures temperatures from +20 on the Holstein Bank without harm. sometimes more than 2 in 25 m. is eury thermal. Under the more usual temperature relations. intergrade. In fact. tropical or subtropical vegetation were free from Greenland and coral reefs could flourish in high latitudes as they are known to have done from fossil remains.. rather appears that temperatures in the polar regions have fluctuated widely. and in general the Salpae and Heteropoda. The origin of the thermocline 15 is explained by the evaporation from the surface. This level is called the "thermocline. so that it sinks to a level where its density is equaled by that of the colder water below. 2 to oyster Ostrea edulis. bivalve Cardium edule and the lugworm Arenicola are others. The thermocline often is the border between two different animal communities. in fact. Examples of stenothermal. which. warmth-limited forms are the reef corals. if attached to The rocks near the tide limit. The temperature relations of the oceans have played an important part in the whole history of the evolution of the marine faunae and of land animals as well." It is found in all warm seas and usually lies between a depth of 50 and 150 m. ranging only between the 20 isotherms of the surface waters. that there have been successive ice ages and successive periods when mild temperatures were world wide and even the polar seas ice the year around. Nothing is thus far known as to what may be the physical basis of adjustments to such variations. The The fective barriers to distribution. the tunicate would grow in Salpa magalhanica. and many other animals belong in this category. the crustacean Copilia mirabilis is confined to temperatures between 23 and 29. which makes the surface water slightly more dense. these temperature relations have not consisted of a simple cooling process that allowed life to originate in the first-cooled waters of the polar seas it and now to spread gradually south as world temperatures fell. Eurythermy and stenothermy. Amphipods tend to be cold-tolerant stenothermal animals. may be exposed to temperatures below freezing at one time and to the direct rays of the summer sun at another. Among individual species. greater temperature differences in the sea obviously form efThus the faunae of the west and east . and even more so the barnacles like Balanus balanoides. and find their principal development in the polar seas. Contrary to the older point of view.196 Physical Conditions in the Ocean notably greater than in the sfrata above and below.

about a third are confined to the west coast.7. the North Sea forms an important temperature barrier. gives the fauna a tropical char- while on the east coast. on the other side of the ridge. at least as far as Cape Canaveral. Similar differences exist between the west sides of Spitzbergen. the much lower temperatures of the eastern coasts ample. coast. and about a fifth to the east. the temperature is about 6 south. 82 The Gulf which washes the west Stream. and of 97 on the south side. or from warm waters 13 to cold. the west coast is reached by outlying of Gulf Stream. The larger size of northern and cold water marine forms is a fre- . Fewer than half the species of mollusks are common to the two sides acter. the arctic ern forms to the southwest.41. Among acanthopterygian fishes the vertebrae number uni- formly 24 in all the tropical species of a whole series of families. and south- Out area only 48. north of it and east the difference between surface and bottom temperatures is 7. while many elongate fishes of the tropics have acquired the same body form by increase in the length of the individual vertebrae. and the character of its fish fauna.8. there is a cold countercurrent that brings the Carolinian fauna southward. (Fig. while to the south it is Atlantic. whereas the cool-water species belonging to these same families have an increased number. 85 are absent on the south. sinae. scarcely a degree further to the 0. The fauna of the northern side of the bank is thus mark- edly boreal. between the Shetland and Faeroe Islands. 4 Finally. 15 fail to reach the northeastern part of the North Sea. or 12. 16). while to the south.-* we may mention Northeast of this ridge the temperature at 1100-m. for exthe parts is Atlantic. Notable among these is the increased number of vertebrae in teleostean fishes from the tropics north.Heat and Temperature coasts 197 of Florida have a very different composition. depth is while at the same depth. 21 The Dogger Bank in bring with them an arctic faunal character. the Among number of vertebrae increases flounders of the subfamily Hippoglosfrom 35 in the more southern to 50 in the Arctic. 14 species on the American coasts Increase in length in arctic fishes is* produced by increase in the number of vertebrae. are common Marine animals present a number of correlations of structure with of 385 species of animals in this to the two sides of the ridge. without increase in their number. the abrupt temperature barrier formed by the submarine Wyville-Thomson Ridge. temperature.5%. Accordingly two entirely distinct faunae are brought into to the northeast of the ridge immediate contact. of the peninsula. where the cold northern water is kept out. out of 167 crustaceans on the north side. the difference is only 0.

but the frequency with which these means confined phenomena apseas inantarctic in and arctic the most diverse groups pear among dicates a convergent association with low temperatures. 16. Delayed favors growth in size. appears to be directly dependent on retardation of growth at lower temperatures. Diagrammatic vertical section across the Wyville-Thomson Ridge with the temperature stratification shown. and this is a common characteristic 11 of northern marine creatures. both as applying to the that replace ranging species and to the northern or cold-water species sexual maturity. with more comthe of embryo within the egg. connecting link between discussion of the physical conditions in the oceans in the present chapter and the chemistry of the sea in the one to follow. fitting Their consideration forms a . 1500 FIG. and the brood care plete development 1000. 11 Water masses The oceans consist of distinct water masses each separated from its neighbors by transition zones rather than by sharp boundaries. The different masses are identifiable by their temperature-salinity characteristics. Large-yolked eggs.198 Physical Conditions in the Ocean individuals of a widequent phenomenon. which tropical forms within a given genus. or ovoviviparity associated with such eggs. are by no to cold waters.

if its arctic extension is considered as north. by Sverdrup. particularly in middle latitudes. torial Unlike the Pacific and Indian oceans. (4) Lower Deeper Water. The subantarctic water mass extends northward in each ocean as Intermediate Water sandwiched according to density between the appropriately named Central Water and the Deep Water below. continental slope. the North Polar Sea comparable to an enlarged Mediterranean or Caribbean sea. arranged in the following order: (1) Subantarctic Upper Water. and diagrams. and large-scale . This whole North Atlantic system of water masses overlies the southward-flowing North Atlantic Deep and Bottom transition Waters. It comes up Water about a hundred meters below the surface of the sea and overlies Antarctic Bottom Water. Both intermediate waters extend over a considerable expanse of more dense Mediterranean Water fanning out from the Strait of Gibraltar. Johnson. The intermediate water mass of subantarctic origin extends as much as 20 north of the equator in the Atlantic but does not cross the equator in the other oceans. and great seasonal variations occur. Temperature-salinity values vary greatly within this layer. South Atlantic Central Water passes by a broad into North Atlantic Central Water that overlies Arctic Intermediate and the transequatorial extension of Antarctic Intermediate Water. (3) Upper Deeper Water.Water Masses 199 Typically the oceans have a relatively thin surface layer about 100 to 200 m. transition masses also exist. To the north. (2) Antarctic Intermediate Water. oceanic current. the Atlantic lacks an equawater mass. The Pacific and Indian oceans are practically closed to the bays. five vertical water masses can usually be distinguished. and so is the Atlantic. Selected examples will be given to convey a general idea of the masses occurring in the oceans and their mode of origin. and Atlantic oceans somewhat resembling great annectant Indian. thick. Water surrounds Antarctica. Ocean extends around the world. and Fleming. The subject is discussed in much detail and with the aid of maps. in the Antarctic Circumpolar to the Surface antiboreal or subantarctic region. and (5) Bottom Water. tables. Each of the larger bodies of water has its own set Antarctic of water masses that despite their basic similarities are organized into The geographic patterns. 34 with particular fairly distinctive No attention to the relation between water masses and ocean currents. attempt will be made here to catalog all the water masses. with the Pacific. Coastal.

] 190 figs. 3:203-214. "Die zoologischen Arbeiten der Kieler Kommission 1870 bis 1920. Akad.s. Chapel Hill. at some season of the year. . Ehlers. ment des mers Blanche." 4. Zool. Coker. figs. wi&s. A. Philadelphia.. 361). 550 figs. the horizontal temperature-salinity relations are similar to the vertical ones of the given mass. of North 7.] 3. W. Brand. 1898. 14: 2 vols... Zool. S. Univ. [Facsimile lithoprint ed. G. Clarke. H. or cold temperate regions. "The action of some denitrifying bacteria in tropical and temperate seas and the bacterial precipitation of calcium carbonate in the sea. Supplement a la faune des decapodes de la rner Ann. S. 9 figs." Cambridge Natural Hist. and along the Atlantic Coast of the United States from 1877-1880. they sink and spread as a result of the relation between their own density and the general distribution of density in the oceans.. Set. Beebe. Schmidt. Ernst. and Karl P.. L." Z. Am. Coast and Geodetic Survey steamer Blake in the Gulf of Mexico. 10:27-38. 8. musee zool. Zoo/. Ann Arbor. Cooke. E." ("Classification of fishes. 2a. Marine Biol. 311 (pp. la. 1949. "Sistema ryboobraznykh i ryb." Festschr. Adv. 1888.. such water masses have their origin in polar. 837. United Kingdom." J. C. by Edwards Bros." 6. Emerson. 1934.. Preuss. Nauk SSSR. Agassiz. 1947. Drew. G. "Molluscs. Caribbean Sea. Saunders. R. Again with two exceptions. (n. Unters. A. "Recherches sur la biologic et zoogeographie principalrusses: IV. Inst. figs. nyne zhivushehikh i [Text in iskopaemykh.200 Physical Conditions in the Ocean Most subsurface water masses are formed when in contact with the atmosphere. Alexander. The principles of animal ecology. St.") Trud. 1875. 9. 1947. Mich. xii. 1895. Meere. p. "Beitrage zur Kenntnis der Verticalverbreitung der Borstenwurmer im Meere. 305. "Three cruises of the U. 5. Orlando Park. The general rule that all oceanic water masses originate at the surface has two exceptions in that the equatorial masses of the Indian and Pacific oceans are formed by subsurface mixing. both recent and fossil. no subsurface water mass is formed at the sea surface in low latitudes. 1921. 325.. L. William. 218). provided by the intermediate masses originating in the Mediterranean and Red seas respectively. Alfred E. Allee. 25:1-102. illus. Museum Camp. 80:495-496. Thomas Park. Assoc. Assoc. 5 (2):1-517. "Oceanographical work at Bermuda of the New York Zoological Society. subpolar. Carolina Press: xvii. Birula.. Karl. rather.. This great and wide sea. BIBLIOGRAPHY 1.) 9:142-155. 1940. (vol. Pub. 1913. 1. Petersbourg. 1939. The place of origin is indicated by the area in which. Berg.. Russian and English. A. 263 2. H. 3:1-459. 1921: 76-194. "The utilization of solar energy by aquatic organisms." Science. Komm.." Butt.

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4:523-529.. Storer. figs. N. "The distribution and conditions of existence of bacteria in the sea. Adolf. 34." 39. U. Teubner: xv. Stimpson. Y. 64: 183-205. ZoBell. McGraw-Hill: xii. "A study of light penetration into photoelectric cell with particular reference to the distribution of plants. Y.. 1946. 5 Naturalist..: xv. Johnson. S. 1943. V. S. and R. 12 figs. 1942. Biol. Steuer. Waltham. The oceans.202 sea water Physical Conditions in the and F. Sverdrup. 1 pi. A. W. 798. Chronica Botan... 1087. Welch. "Distribution marine shells of Florida." 31.. 4:586-587. made with the Kung 1922. I. pis. chemistry. A. text figs. E. Shelford.. S. Corey.. Waksman. 265 35. T. W. 1910. Ocean 30. William. and general biology. Puget Sound Biol Sta. 551 figs.. Prentice-Hall: x.. Gail. Renn. 723. 3:141-176. M. "Microbiological activities at low temperatures with Quart.. Pub.. Bacteriological investigations of the sea water and marine bottoms. General zoology. H. 38. 7 charts. Monographs. H. 365 of the 32. Idem. E. L. 471." 37." Bull. Limnology. Rev. McGraw-Hill: Ecol. C.. III. 1934. and C. 9:460-466. M. 1871. . N. Hotchkiss. 1935. xiv. particular reference to marine bacteria. C.. Marine microbiology. 240. 46 figs. "Studies on the biology and chemistry of the Gulf of Maine: Biol. a monograph on hydrobacteriology. their physics. W. P. Reuszer. N. Leipzig. 33. E. Y. 1933. 36. H. 1934. Waksman. Planktonkunde." Am. Fleming.

6'^. in the western Baltic the salt content is 12'/. where a bottom current of more concentrated salt water flows in while the more dilute surface water flows out. the solution melting regions The most marked deviations from the average composition occur in where exactly the same. 39.5 /{ f salt. at Crete. The fresh water Bornholm. By contrast. and iodinemake up about 55% of the dissolved solids. The intermixture of their waters with the open is limited by their degree of separation.5' vr. its relatively low influx of fresh water. Their quantitative analysis is readily 203 .. and differences in one Thus the Mediterranean. "Halinity" would be a preferable term. ( has as much as 45. with no regular supply of fresh water. and in the Gulf of from melting glaciers and ice- may amount to "Chlorinity" is often used in determining salinity or even as a working substitute for it. In the polar the ice causes a dilution of in summer. bergs dilutes the surface water of the polar seas so that the salt conc As the more concentrated solution is f o. lA'/co at Finland only 0... The Chemistry of the Sea Its and Salt content Influence on Animal Life SEA WATER IS A SOLUTION THE COMPOSITION OF WHICH IS NOT EVERY- In the open ocean the salt content at 300-m. thus in the western Baltic the salinity at the surface 8-12'/r and in the depths may reach 27'/tr>. In the tropics the evaporation exceeds the salt solution becomes more concentrated. The Red Sea. but there are differences in the composition of the surface water. depth approximates 35'/e.. the salt content may increase with depth. and the ocean with the seas and gulfs. has a salt content of more its than ST/tc in western part. This is especially evident in the Baltic. bromine.II. direction or the other arise accordingly. The halogen ionschlorine. and even 40% on the Syrian coast. tent may fall below 15 / denser. inflow.4^<> to 46.

magnesium. like silver. greater part of the salts in sea water consists of the chlorides of sodium. are adapted to the medium in which they live in having their . which predominate in the sea water. The with 27. Osmotic pressure rises with increasing concentration and It amounts to no less than 26. which use strontium sulphate. Indianimals profit by the substances taken up directly by the rectly. as. and draw calcium. Other elements are present in much smaller amounts.9 atmospheres. and also silicic acid. of 7. magnesium. but a condition that does not concern such animals in nature. for example. they are usually normal sea slightly less than 19.204 The Chemistry titration of the Sea made by water is with silver nitrate. calcium. the radiolarians of the suborder Acantharia. may contain Epsom the composition salts or borax. Seas without outlet salinity is exclusively derived from river water do not have of the sea water." and the chlorinity of so-called Combined. at 18. is formerly less saline than at present. Animals are able to directly upon a number of the inorganic materials of the sea water to build up their bodies. horn corals store bromine and iodine in their skeletons. referred to as "chlorine. plants. for example. copper. especially sodium. of which common salt (NaCl). furthermore.5%c salinity. of 40%o salinity. and ammonia. sulphur. Salt masses from itself may have conditioned the composition of the the sea are important in the life of its animals. sodium. To these must be added the sulphates of magnesium and calcium.7 falls with decreasing concentration. and vanadium. in addition to potassium. nitrates. and. Rivers. The assumption that the sea salt derived from the land is not necessarily true. per liter. only The question as to whether the salts of the sea water have been derived from the earth by the solvent action of the rains and streams touches us in this connection only in so far as it bears on the biologically not unimportant problem as to whether the sea was in traces. nitrites. which are present in excess. Occasional animals may use rarer elements. contain only very small amounts of chlorides. and chlorine. even when this is accomplished gradually. with the exception of the bony fishes. it is only 4. plants use among others The salts of phosphoric acid. The differential it withdrawal of NaCl is as a coloring mateis fatal to most marine animals. some. at 30. The osmotic properties of sea water are of high importance to marine life.37 gm. rial in their and ascidians contain vanadium blood. in the of water the Red Sea. atmospheres In the waters of the Baltic at Bornholm. whose the sea bottom sea water.4?. magnesium. and potassium. Marine animals. preponderates.

reef corals. of course. the inhabitants whose waters are concentrated by the summer sun and freshened by fall rains and melting snow in the spring. stenohaline animals. where the ever. stenohaline Euryhaline forms include the coastal pursuit by fauna that lives between the tide levels and is thus exposed to rain water at low tide. The former animals are characterized as stenohaline. able to live side by side with stenohaline forms. Examples of euryhaline animals are such jellyfish as Aurelia aurita and Crambessa tagi y which are driven into river mouths without harm. stages of the fish Fundulus. There are all possible degrees of the sensibility to variation of salt content. so that these are not altered by probably for this reason that some marine animals are very sensitive to variations in the salinity of the surrounding water. Closely related animals behave differAmong the chaetognathous worms. limits of variation in this respect. the animals near the mouths of rivers. and the fauna of spray-pools on rock coasts. extent of the influence of the fresh water varies with the variable volume of flow of the of the salt marshes. with sea water. Others. and in fact . for which depth rains may dilute the sea water. river and the state of the tide. whereas Sagitta bipunctata 24 has an unquestionable adaptibility to brackish water. the latter as euryhaline. where they are little exposed to this variation. lugworm Arenicola. as in saline lakes of the faunae of waters with varying salinity is and in the Baltic. They are found primarily in the open ocean. are accessible only to them. some sharks that range into fresh the water. can escape this variation at the of a few meters. Stenohaline animals are subject to certain restrictions in their distribution. and vice versa. on the contrary. the edible mussel Mytilus. above all from freedom advantage organisms enemies.Salt Content 205 body fluids isotonic It is diffusion. Stenohaline animals may live on the coasts tidal variation. and some can even withstand relatively rapid change from weak to strong concentration. hexaptera is very sensitive to fresh water. away from river mouths and below the level of are also suitable for them. the green crab Carcinides maenas. Great depths Colonial radiolarians. are able to live in waters of widely different salinities. juvenile and many others afford examples of Euryhaline animals are. the appendiculate Oikopleura dioica. and numerous others. in habitats in which they make no use of their Other situations. Sagitta The composition regulated by selection. and more exclusively so with greater Such situations afford euryhaline the of from competition. ently in this respect. howability to withstand variation of salt content. Even the surface forms.

which is also able to resist highly concentrated saline size. Among fishes the Syngnathidae are inclined No group of animals has produced as many Whole genera of lamellihabitat. so that the brackish-water derivative remains as a relict. the latter six. Brachiopoda. so that the direct relation with their marine ancestor is no longer evident. in length from the North Sea and the Baltic. branchs and snails have gone over to this ber of brackish-water forms occur at present where the immigration from the sea into this habitat and into fresh water appears to be much easier than in higher latitudes. but do not reach more than a third of the normal size. to take to brackish water. The small number of stenohaline brackish-water animals con- fined to brackish waters includes the hydroid polyp Cordylophora lacustris and crustaceans such as Eurytemora hirundo and Temora longicornis of the Baltic.206 The Chemistry of the Sea in all brackish waters. are entirely absent. as well as Cephalopoda and Selachia. This is also plainly shown in the impoverished fauna of the Caspian Sea. The greater numin the tropics. degree. it does not retain calcium carsalts It bonate in solution. its marine ancestors have been found fossil in the Miocene of North America and the Argentine. or by the extinction of the ancestral form in the marine habitat.59 to 1. 13 Calcium compounds Calcium is the only one among the more important constituents of the sea whose proportionate amount fluctuates to a significant varies from 1. important groups of animals. Adaptation to lowered salinity is frequently connected with reduced Sea urchins wander into the mouth of the Rhone as larvae and become mature. averaging 1. Scaphopoda. scyphozoan medusae. 17 Marine bivalves exhibit salt strikingly with reduced content of the water. brackish-water forms as the mollusks.6%. The latter seems to be plainly the case with the clam Tagelus gibbus. Echinodermata. and yet there can be no doubt of the marine origin of the fauna. Ctenophora. widespread in the ocean. Tunicata. such as Anthozoa. Among the most persistent of the inhabitants of the brackish waters of the Baltic and of the Mediterranean lagoons is the annelid Nereis diversicolor. the former was three years old. Pycnogonida. reduced size in marine basins Heincke compares two flounders (Pleuronectes platessa) 21 cm. 10 waters. . now confined to brackish waters in America. and the calcium occurs principally as sulphate. the reasons for this are still unknown. The production of special brackish-water forms may occur in two ways: by mutation. Since sea water is in general alkaline.82$ of the total salts.

by (NH 4 )L>CO 3 to a considerable extent by temperature. the calcium in the sea water is inaccessible to the marine animals. serpulids. Calcium deposits by animal life reach their maximum in the tropics. CaSO 4 is usually precipitated for this purpose Experiments show that this reaction proceeds very 21 rapidly at 26-29. for example. In very cold waters. of lime are also absent in deep seas. The free use of calcium salts by marine animals is thus conditioned armor. as its shell lengthens with increasing age. Calcium is utilized by innumerable animals in building up spicules. The serpulid worms.Murray has pointed out that the calcium. without shell or with a delicate chitinous armor. which build lime tubes. It observation that crabs change their shells has been shown by direct and grow new ones more Frc. and Pteropoda. rapidly at high temperatures than at low. but very slowly at lower temperatures (4-7 ). the snail Magilus. . as among Coccolithophoridae. Magilus antiquus with the coral limestone cut away to show the which is filled with calcium carbonate to the dotted line. shells. which is able to dissolve the carbonate. and skeletons. and the tropical forms with strong shells or skeletons of this material are accordingly often replaced in polar seas by naked or soft forms. also reach their highest development in tropical seas and have few Animals that deposit large amounts representatives in polar waters. where the reef corals build gigantic masses of rock. shell. Foraminifera. of The shells of the giant clam Tridacna gigas may reach a weight more than 250 kg. 17. fills a large part of it with lime. thanks to the presence of free carbon dioxide. 17).Calcium Compounds in contrast 207 with the condition in fresh water. . ing. are want- and the deep-sea sea urchins are forms with a soft shell This constitutes an important factor in the distribuEchinothuridae ) ( 10 2 tion of marine animals. living among coral reefs (Fig. . Tropical mollusks often have shells of great thickness.

14 in December pH 8. accordingly flourish most where there is a constant inflow of water with available carbon dioxide. The presence of the inorganic substances of primary importance to plant life. and partly by the large amounts constantly brought in by the rivers. more and more in the tropics at the present time. so that carbon dioxide is stored in the depths. determines the quantitative distribution of plant life in the sea and thus secondarily influences the distribution of animal life. per cu. together with carbon dioxide and phosphoric acid. Nitrogen in combination as nitrates. phosphoric acid. The observations of Atkins 2 show that the carbon dioxide in the upper layers may be rapidly used up. The amount in milligrams varies with the temperature. Nitrogen compounds similar with the nitrogen compounds. is present in the North Sea to the amount of about 300 mg. This store becomes available to marine plants only where vertical currents bring the bottom water to the surface. and the decomposition of the sinking bodies of plants and animals adds to this. and rather less in the waters of the open ocean. carbon dioxide. carbon dioxide per liter. is The expended carbon dioxide that from the The animals inhabiting the aphotic depths give off strata of water. the excess of basic compounds. Sea water contains 40 to 50 mg. 4 which are unconditionally necessary to plant life. Carbon dioxide is used by plants only in the euphotic and dysphotic bonates. He found that the reaction of the surface water on to the English coast varied with the season from pll 8. and the nitrogen compounds. and thus available to plants. on account of solution. such as carbon dioxide. nitrites. m. This seemingly small variation really amounts to a fall of 25% in the hydrogen-ion concentration and results from the accumulation of the carbon dioxide produced by the excess of animals in winter and its Marine plants use by plants in their flourishing summer growth. Experiments have made it certain that plants are able to use the carbon dioxide contained in bicar- replaced in the sea partly air. The situation is or ammonia. about as follows: .27 in May at the time of the highest amount of sunshine. partly by produced by the marine animals.. of which only a few tenths of a cubic centimeter per liter is present in simple it is combined as carbonate or bicarbonate.208 which the The Chemistry rivers bring into the is of the Sea ocean and which comes from the thus deposited decomposition of continental rocks. especially those not present in excess.

on the average. the nitrogen is available for green plants only in the lighted upper levels. and by rains. cited worth rereading.8 27. it seems likely that the smallness of the amounts present in sea water limits the amount of plant life and thus also the animal life of a given part in the sea are various. The different amounts of these fertilizers in different areas explains the difference in the abundance of marine life. are compounds by the considerable "ammonia pressure. and the inflow of fresh water. plant life will At greater depths (5400 m. general mixture of marine water to the bottom occurs only in A . the upwelling of deep water.7 N as N2O3 and N2O6 N as NH 3 210 68 144 142 91 75 46 36 78 45 In view of the great requirements of plants for nitrogen. at 400 m. of the sea. fact that the bodies of many dead organisms sink far below the plant- inhabited levels before they decompose.2 12. and ammonia. The question whether nitrifying bacteria normally occur in sea water has not been Waksman and of ZoBell. where it is absorbed by rain drops and transmitted to the soil. 313 mg. at 800 m." passing into the atmosphere and blowing over the land. also in organic compounds. 1 Where such bottom waters The fertilization of the lighted upper levels of the ocean by nitrog- enous compounds thus occurs in a variety of ways.Nitrogen Compounds Average temperature 3. Brandt ) The loss of nitrogen in the upper layers is increased by the ( . per cu. as for carbon dioxide.4 209 23. principally through the thorough mixing of the waters. nitrogen present was approximately constant from surface to bottom " There is conflicting evidence on this matter. which is freed from the ammonium settled with finality. m.. and 485 mg. The sources of nitrogenous compounds They originate in the seas themselves by the metabolism of animals and by the decay of dead organisms.0 17. The supplies are diminished by loss of ammonia. and perhaps which contain a little nitric acid. The supply from without is brought in by rivers in the form of nitrites. nitrogen as nitrate and nitrite occurs in the oceanic surface water to the amount of 101 mg. Nitrate-fixing bacteria may well be present on the ocean bottom. Thus in moderate depths.. are brought to the surface. water near the bottom of the sea is thought to be richer in nitrogen. In any case. The nitrogen content consequently also increases with depth. especially in the zones of frequent electric storms.) Krogh found that the total flourish. at the time tested. nitrates. The conclusions of ii the introduction to this section.

There is a periodical mixing of the waters. the guano deposits of the outlying Peruvian islands are the indirect and the luxuriance of the submarine forests of algae (Macrocystis pyrifera) with their rich fauna. the greater concentration of organisms in the milder part of the Arctic and Antarctic seas. with its renowned sardine fisheries. south of Iceland. It is The Chemistry produced in of the Sea shallow coastal waters by the tides. to greater depths by storms. where continued offshore winds carry away the surface a water. as in the Baltic. The mix- ing extends to a depth of 140 m.210 shallow seas. and the "red tide" of the Japanese coasts. the west coast of Morocco and the African coast opposite the Canary and Cape Verde Islands. Their origin is betrayed by the coldness of the water. in consequence of the cooling of the surface water in the winter and season. Coloring of the sea by thick masses of plankton appears only in shallow seas. No oceanic waters swarm with so much life as the upward streams of deep water in Such regions are consequently tropical and subtropical latitudes. doubtless also depends primarily on this kind of fertilization. to 800 or 900 m. which are rich in nitrogenous compounds. 7 The Kiel Fiord exhibits an occasional upward current of cold water. It also explains. some places on the Portuguese coast. and shallow banks. in the Atlantic and to greater depths in cold seas. Such currents stir up the bottom strata. have such a rich flora and in consequence a rich fauna. such as the Dogger and Newfoundland banks. . in part. and coasts On where they become available as foodstuffs. and the coast of southwest Africa. the Gulf of California ("Purple Sea"). The deeper the ocean the more difficult such mixing of the sea water from the surface to bottom becomes. and thus become favored fishing grounds. in the temperate and cold zones. The coasts of Chile and Peru. carry them to the surface. A southwest Or west wind drives the surface water out of this bay so that the bottom waters rise. The is astonishing richness of the fish fauna in the cold-water areas of the Arabian Sea near Oman and at Cape Hafun on the Somali Coast even more striking. reasons why and why shallow seas. fisheries. and this is sufficient to bring up the nitrogenous water from the bottom in shallow seas. like the North Sea and the western Baltic. Such localities include the Algerian extensive sea to adapted coast. when the westerly wind prevails result. on the coast of Chile. are especially notable for wealth of marine life. the Gulf of Guinea. with their cold 3 upward currents. compensating current rises from the depths. This is one of the coastal waters in general are richer in animal life than the near-by deeper seas with similar physical conditions (light and heat).

of phosphorus for nitrogen. pass the west coast of Japan where the is Deep water drawn to the surface even when two neighboring cur- rents have a similar temperature if the currents are diverted in opposite directions. Both localities are the sites of important fisheries. whose waters are no colder than 12. are poor in life. north of Ascension Island.Phosphates for a considerable time. as by the Wyville-Thomson Ridge. accompanied by seasonal Phosphates variations in the quantity of plankton organisms. in part because they are presThe ratio of marine nitrate-nitrogen ent in smaller concentration. Examples are presented by the Newfoundland Banks. Both have seasonal to variations with lower concentrations during the "regeneration" of phosphates summer growing and is season. The small importance of thermal currents in the Mediterranean.9 at the bottom. where the surface water is also rich in life. and warm Kuroshio and the cold Ojashio each other. Downward currents attract surface water whose nitrogen and carbon dioxide have been exhausted. Phosphates may be the more important. and areas where they occur. or 7:1 in weight. the 211 water of the fiord become rents 5 grass green from the multitude of diatoms. with its richness may be ascribed to this cause. in consequence of the differing densities of the adjacent masses of water. The development of diatoms may with northerly and easterly winds. Many of the relations and silicon are similar to those that have been discussed ations of the amounts of these In the higher latitudes there are distinct seasonal fluctuessential mineral nutrients. phosphate-phosphorus in the seas roughly approaches 15:1 in milligram-atoms. The takes less time . The existence of the so-called "tongue of cold" of the South Equatorial Current. as in early spring. and in the polar seas the deepest waters are involved in the movement. The Straits of Messina between Italy and Sicily also have a rich plankton. Vertical mixing also appears in places where a warm current passes a cold one. The two sets of chemicals are similarly distributed both regionally and in depth. like the Sargasso Sea. Deep-sea curbe diverted into an upward direction by submarine banks is slight and ridges. Phosphates in the sea resemble nitrogen compounds in their importance as limiting factors in the development of plant plankton. where the Labrador Current and the Gulf Stream adjoin. mixed in consequence of the different tidal phases of the Tyrrhenian and Ionian seas to the north. south of the Faeroes. the waters are in plankton. results in general poverty in plankton despite local exceptions.

are carried toward the coast of Greenland by a westward curof river water. contributes to the same results. The Atlantic has in rent..212 apparently less The Chemistry complicated than is of the Sea 11 the similar release of nitrates. about three-quarters of the observed phosphorus near Bay of Maine comes from the depths by vertical transport.000. The Atlantic-Arctic is thus surrounded by tributary land masses. ) . km. Influence of rivers present in trace concentrations only. are brought to the sea in large amounts by river drainage from fertile areas. life in coastal waters. the major part of North America and almost the whole of South America drain into the Atlantic. from the Pacific Ocean biologically as well as geographically. though of the entire Pacific Ocean is more than 3600 m. eastern Pacific is open southeastern that must be added almost three-fourths of the extent Pacific. many 25 in the economy of the sea. with 265. and with a surface of 103. The surface water contains.. are also important The distribution of rich faunae in the ocean is influenced also by the amount of materials received from the land. Phosphates. receives the drainage from over half the land surface of the earth. per cubic meter ( the figures : in parentheses refer to the tropical belt between 24 S. contrary. Compounds of silicon.000. receives its land waters almost entirely from Asia. in the oceans. especially at the north. and the waters of the mighty Siberian rivers. 25 The is least supplied with land waters. principally from the This is probably an additional factor in the wealth of marine rivers. carbon.. and other chemical ele- ments. with it These facts help explain the poverty in life of the the lack of coastal habitat differentiation. and 25 N.000 sq. and returns from its enriched on the Indo-Pacific-Antarctic. with 13% of its extent within the 200-m. deep and that the absence of a continental shelf on the west coast of the Americas. line and only 47% deeper than 400 m. and this source of almost The result is that the Atlantic differs fertility negligible. The southso is well equipped to produce maximum waters. The waters of the Pacific have the lowest content in nitrogen compounds. it receives the drainage of only 27% of the land surface. but even with ready regeneration and the surface of the 14 inflow.000 sq. addition wide shallow areas. draining into the Arctic. Such materials are very unequally distributed The Atlantic and Arctic receive the largest amounts They receive the waters of all the European rivers and of the most important African rivers. km. like nitrates.

(43 mg. The distribution of life in the sea depends on the fertility of the water. Alexander Agassiz l refers to the middle Pacific as "barren grounds. may be many times less than in good soil. on account of the presence of of oxygen at salts.) 119 mg. algae. The amount of gas absorbed by water is larger at low than at high temperatures.) 52 mg. and this necessitates that they of small size in order to have a high ratio between area of surface be and bulk. (50 mg. of the Chinese coast are among The rivers of southern Asia <J Even at its best. and grasses of the land. The general conclusion that the tropical seas are poor and the cooler and cold seas rich in plankton. sea water is a very dilute solution of many of the mineral nutrients essential for plant growth. The western Pacific. Chun finds the crystal-clear surface waters between the Seychelles and the east African coast poor in animal life. at and only 4. The tropical parts of the oceans are probably poorest in the only exceptions being places where there is extensive upwelling of the bottom waters. In contrast to the trees. even of essential substances. shrubs.) life. 25. and the amount dissolved in sea water is accordingly greater at the poles than at the equator.) 46 mg. (66 mg.Oxygen Ocean Atlantic 213 Nitrogen in Nitrogen in Nitrates 102 mg. is much richer in life. even smaller. and this depends in turn on a number of factors varying from place to place and sometimes from season to season. (46 mg. requires some modification.) Ammonia Indian Pacific 47 mg. into which the great Asiatic rivers flow.03 cc. although relatively true. (88 mg. Mitsukuri speaks of the plankton of the Japanese sea between Honshu and Shikoku as quantitatively the richest known to him. the marine plants that form the base of the food chain consist of minute diatoms and other. the concentration.93 salinity may contain 8. A liter of sea water of 35# cc. and the great extent of the coast lines in the East Indies are favorable to a rich marine life. together with bacteria. The fisheries of the Japanese coast and of part the most productive in the world. with its wide shallow areas and good development of coastal 18 formations." in sharp contrast with the abundance of life in the Gulf of Mexico or in the main current of the Gulf Stream along the Florida Reef. oxygen is one of the most important for marine life. In contrast. oxygen . This means that planktonic plants must secure salts many of their essential from an extremely dilute solution. Oxygen Of the substances dissolved in sea water. (77 mg.) 69 mg.

. The most superused oxygen from the air. there may be a deficiency of almost 100$. Within the lighted zone the metabolism of plants is an important source of oxygen. like the worm Spio juliginosus in the Bay of Naples. up to 9. 15 Hydrogen sulphide also accumu- . Bottom life becomes scarce in such places. K In the deep troughs of the Baltic the water stagnates. Bqrnholm deeps. Natterer found the bottom waters at 3700 m. The depths of the Black Sea contain intubicolous creasing amounts of this poisonous gas below 180 m. deeper waters of the Atlantic have a deficiency of only 25%. but even 2. In the lower about ficial liberate. In many places where there is stagnation of the water at the bottom but a rich surface fauna. Thus in the Atlantic at 500 m.60 cc. which receive their waters from high When latitudes. all life is absent. in latitude 10 N. and harbors accumulate a black ooze filled with hydrogen sulphide. of oxygen per liter will support much life. in a In the Mogilnoje Sea (Murman coast) the hydrogen sulphide exliter." In the Gulf of Maine this compensation point is mostly around a depth of 25 m. In the great oceanic depths. or is reaches of the lighted zone. 4. with only 2.64 cc. plants consume as much oxygen as they more. carbon dioxide accumulates bottom and there is a lack of oxygen. there is frequently greater oxygen deficiency than at lower depths. the level at which consumption balances production called the "compensation point. with a resultant lack of oxygen in the bottom waters and with attendant reduction in the number and of species at these places.. The Bay of Naples and the Gulf of Guinea on the west African coast are examples. The bottoms of isolated bays. This occurs in some places in the Mediterranean.214 is The Chemistry 208? less soluble in sea of the Sea than in fresh water. and it is distributed to layers replace moderate depths by wave action. lagoons. northern Balearic Sea) which extend there is at the to great depths.. the want of oxygen becomes so great that hydrogen sulphide produced by the decomposition of animal bodies is no longer oxidized and accumulates in the deeper water. 9 In depths of about 500 m. tends to within 13 m.. with the exception of anaerobic forms.58 cc. 3. thus affording a limited oxygen supply.64 cc. descending currents of surface water in the Mediterranean at isolated places (Aegean Sea. Where the accumulation of hydrogen sulphide continues. at another place at 1005 m. southern Adriatic. There are.. the Gotland deeps 23 are such localities. the oxygen content is only a little below normal. and only the most resistant forms persist. no such bottom drift of cold water. of oxygen per liter. Bay of Danzig. of the surface. and at a third at 1210 m.85 cc. however.

" Part Marine Biol Assoc. 166-206.. 4. R.. 551.. and in the Gulf of California. Teil 2." pp. figs.) connected with the sea by shallow channels. Beagle under the command of Capt. London. Zool. J. Jena. 12 3. 13:93-118. R. 20 A number Gulf of Guinea of reports of great mortality in the fishes of the are on record. Atkins. chemistry 1-194. Agassiz.N. 1935. C. chemistry and turbidity. Journal of researches into the natural history (p. 9.. Meeresunters. Fitz Roy. illus. 1923. 3. in Bronns Klassen und Orclnungen. 1890. Meeresunters Kiel 4:213-230. Mediterranee. illus. 467-678. "Die Rev." when the cover of ice prevents the oxygenation of the water. to the west coast Mexico. Fcstschr. 3. (p. "Ueber den Stoffwechsel im Meere. Wiss. Drechsel. 1928. Alexander. When storms or occasional advances of cur- rents carry the poisonous gas into the upper levels. Fischer: vi. saltwater basins of moderate depth (up to 30 m. 1891. "Die zoologische Arbeiten der Kieler Kommission 1870 bis 1920. 8:286-287.." (p. "Biological Cambridge.5-79.Bibliography lates in 215 summer at the bottom of certain of the Norwegian fiords. Brandt. Austernbassins in Norwegen. 12 BIBLIOGRAPHY 1. 1:553-573. the animals there may be killed. 6:2." Intern.. Preuss. H. of the countries visited during the voyage and geology round the world of H. R." Kiel: 76-194. 18:185-429. of 2. Cambridge Univ. W. . Board Con. Abt. 390 text figs. 239). and physics of sea water.. Darwin. 9th Intern. 3: 467-678. Carl. 46 pis. Komm." 5.. UydrobioL. Murray: xvi. United Kingdom. Chun. Karl. 1914. C." 1:279-467. 8. 1908. 1899. the hydrogen sulphide may reach the uppermost levels and spoil oysters kept there in baskets." Natunvissenschaften. Press: Holland-Hanson. but in these it is removed by the winter overturn of the waters. 12. "Tiergeographische Strandbeobachtungen von der Westkiiste Siidamerikas. Monaco: 123-219. Biol." H. R. Fritz. F. 1920. In the Norwegian "Polls. 6.. "Three letters on the dredging operations off the west coast of Central America to the Galapagos Islands. 280). B. Brand. and continuations. 594. 1916. 570). 11 figs. Schilderungen von der Deutfschen Tiefsee-Expedition. W.. Bivahia. "A quantitative study of the phytoplankton in the Bay of Fundy and the Gulf of Maine. ion concentration of sea water in II. Zool Congr. H. Aus den Tiefen des Weltmeeres. Harvey. 1941. 11. "The hydrogen figs. Ifg. 1900. 2 maps. 1902." 21:185-200. 10.S. 7. In the summer of 1911 the entire Mytilus culture at Ganzini near Messina died out because the ooze was stirred up by a current. Idem.. Bull. Bjorn.M. Gran. G. including observa/. "Oekologie der Muscheln. tions on hydrography. its relation to photosynthetic changes. 1921. Haas. and Trygve Braarud. Museum Comp. "Les expeditions oceanographiques danoises dans la C.

Entwurf 23. figs. 1912.. John. Acad. Archhelenis und Archinotis gesammelte Beitrdge zur Geschichte der neotropischen Region. (p. N.. 51-53. 19. Zool. R. The N. Hansen. Ketchum.). Dez. and Robert Irvine. Julius. Edinburgh." Ecol. Rudolf von. Ritter-Zahony. 1896." Butt. B. Martin W. Y. illus. HellandAtlantic. Appelof. Soc. 17:309-315. and general biology. August.." Vcrhandl. 2. M.. their physics. The Chemistry of the Sea Hermann von. chemistry. Sci. Sverdrup. "The biochemical relations between marine organisms and their environment. 1898. 14. London. 1 map (p. 16. Macmillan: xx. "Die macroscopischc Fauna dcs Golfes von Neapcl. 21). "Conditions of life in the ocean. Leipzig. 24:221-235 (p. 17. "Distribution geographique. U." C. Plankton Exped. Roy. 4:421-429. and B. Zool. 1911. 7 charts. 1947. and Richard H. 1895. H. Knipowitsch. 4th Intern. 258)." Ergeb. and Johannes Hjort. sci. Kakichi. "Die Bodenfauna von Nord-und Ostsee.. John.216 13. of marine organisms. 1907. 24. St. Murray. 3:459-473. einer biologischen Analyse des Meerwassers. Wilhelmi. Monographs. Priifungsanst. 25. Fleming... H. H. Proc." Zool. 16:47-166. Reibisch. Mitsukuri. 821. "Ueber den Reliktensee 'Mogilnoje* auf der Insel Kildin an der Murman-Kuste. Marion. x." serversorgung Berlin. Murray. 1885. 233). (p. Petersbourg. Engelmann: 350." Ecol. 320 ff.. dent. 3rd Intern. 18. vom Standpunkte der biologischen Analyse des Wassers betrachtet. 109). "On coral reefs and other carbonate of lime formations in modern seas. Congr." Annee biol. J. 21. Gran. Pruvot. Cambridge: 101-111 (p.. 1 fig... 1895. Prentice-Hall: oceans. 26. G. 1912. Monographs. Acad." Proc. 1087.. 2:558-587. Murray. 1934. 15. 1891. lettres beaux arts Marseilles [fide Hesse. John. 22. "Die Chatognathen der Plankton-Expedition. 1942. Johnson. Krogh. H. Ihering. "Zoological matters in Japan. 265 figs. Was- . Heft He: 1-33. Teil. "The general conditions of existence and distribution 20. The depths of the ocean: a general account of the modern science of oceanography baaed largely on the scientific researches of the Norwegian steamer Michael Sars in the North With contributions by A. 17:79-109. 11 figs. Leyden: 99-111. 1914. 1924]. Congr. 1 map. Ges. Mitt.

where conditions approach the optimum. in the coastal areas of warm seas. bear the stamp of a similarity produced by adaptive resemblances in structure and habit. In consequence. according to the steepness of the coast and the 217 . two zoneTmay be set at the bor3er^D>rthe continental shelf. and The boundary between the the coast drops steeply off to great depths. and in brackish seas. in spite of being composed of animals from groups separated in zoological classification. Two subdivisions of the littoral are to be recognized. the animal communities exposed to a given set of conditions. in the deep sea. Such common characters appear more plainly the more numerous the adaptations required by the environment. which on the average lies at a depth of 200 m. In the ocean as a whole.The Biotic Divisions of the the Ocean: Benthal THE VARIETY IN THE ENVIRONMENTAL CONDITIONS IN DIFFERENT PARTS OF the ocean produces corresponding differences in adaptation of the inhabitants. and these become more sharply defined the more extreme the conditions. the possibilities for variety of organization are greatest. as in the littoral area in polar seas. The lighted zone in such places must still be called the littoral. At many places there is no sloping littoral shelf. two is diminished and the variety of main divisions may be contrasted. Every change in the environment in the all direction of the unfavorable brings limitations with it. the rbenthal" and the "pelagial" ( Fig. At the hypothetical original site of the development of life. The lighted zone of the benthal is designated as littoral. 7 18 J/ Each falls vertically into two parts. the sea bottom and the open sea. ajighted zone and a lightless one. At such places the number of species structural types is reduced. the lightless as abyssal.

for example. currents. The borderline between the animals of tide pelagial and benthal is thus not very If one emphasizes the characteristic of free sharp. The benthal The primary difference between the life of the benthal and that of the pelagial lies in the necessity for animals in the latter to be independent of any support except that of the water.218 The Benthal nature of the shore: depositing shore. and quiet water. Primary subdivisions of the marine biocycle. even though they may be able to swim about for a longer or shorter time. swimming too . Johnson. like the scallops among bivalves Qjjjjke the flounders among fishes. however. coral coast. pebble beach. the lighted zone. In the pelagial. which " . are not able to float continuously.. regions of upwelling or sinking waters. sand beach. rock coast. usually with more or less loose shore material. an animal that is capable of remaining suspended in the open water may also move or rest on the bottom. Modified from Sverdrup. The bottom-dwelling forms. 1000 EULITTORAL (SUBLITTORAL ARCH BENTHAL I [ABYSSAL- BENTHAL FIG. Special biotopes in the pelagial are conditioned by the movements of the vvater. and contrasts with the oceanic. and Fleming. etc. mud beach. and eroding shore. Each of these groups divides into a number of biotopes according to the nature of the shore. which lies above the littoral. 18. is designated as the neritic region. more or less steep and rocky.jirir:unnnnrLnnnnn: NERITIC hjrrif^'ijirirv'LririririrLri^ -"Tt" a ir--~_-_-_r-_-_-_-_-_--H. lies over the greater depths and includes them. e. To be sure.g.

littoral benthal. although their food is taken from the open water. while that of motion is subject to various irregularities and special conditions imposed by cosmic and terrestrial influences. the mullet Mullus barbatus beit depends on this section for its food. that of the open water is none the less bottom may be called. extending to the limit of more abundant vegetation at 40 to 60 m. One might hesitate about the classification of the skillfully swimming labrid fishes. the distinction between the bottom fauna and a natural one. "benthos.Subdivision of the Benthal 219 much. Julis. which watch over the eggs that are placed on the bottom. sand.. Within the benthal the different kinds of shore material produce The substratum may be important differences in the environmental factors influencing its life. and the lightless arcbibenthal ancL abyssal benthal." as the animal life of the The although longs to it it because can swim for a long time. In quieter water. Both light and motion diminish with depth. Loose bottom may be coarse or fine gravel. Subdivision of the benthal The sea bottom is accordingly divided vertically into two main zones: the lighted sea bottom. loose or solid. Even though the same species of animal may at some times be pelagic pelagic. more briefly. is of animals or for the most composed part dependent on the wholly bottom for support. line. or mud. occasionally at depths up to 1000 m.) * Carls. We follow The Oceans 82 in recogniz- . Solid shore material is found only on steep slopes and in actively moving water. since and at others benthic. or eulittoral. even fishes like the it is mackerel could be regarded as only partially highly probable that they rest from December to February on the sea bottom after the strain of continued swimming. theTittoral proper. littoral zone may be dividecf into two subdivisions.* or Labrus males. and the sublittoral. which drops off toward the abyssal depths at about the 100-fathom (200-m. from this point to the edge of the continental shelf. Other differences in the life of the benthal depend largely on the motion of the water and on lighting. consequently mostly near the surface. Crenilabrus. but the decrease in light in clear water is uniform. In water subject to motion. Thus the squid Sepia belongs to the benthos. with every sort of transition. such a bottom moves with the water above it. which frequently rest on the bottom to sleep. sediments accumulate and maintain themselves even in the face of slow movement of the water. the The littoral. or.

) above the abyssal Thermal differences appear to offer the best basis for such ing an upper archibenthic zone benthal.6 m. The second zone is that of wave motion. A and are exposed to a special set of conditions. on the other hand. its inhabitants are a mixture of terrestrial and marine animals. but still wave-affected. so that many groups ing forms. with their fixed ancestral stage. animals from just below the low tide mark differ significantly from those of the deeper. The faunae of the deep sea as well the open ocean are nothing gill-breathing as the life of specializations derived from the Bottom life at moderlittoral bottom faunae by special adaptations. and below that is the third zone of quiet water. I Marine animal life is most richly developed both in number of . bounded above and below by high. in the Bay of Fundy.220 The Benthal ( 200 m. This is especially evident in the pelagic medusae. are plainly derived forms. the difference amounts to 11. Nova Scotia. and the pelagic mollusks. ate depths in coastal waters requires the fewest adaptations to physical all groups of marine animals are represented. to 1000 m. 12. Normandy. waters.and low-tide marks. thus at Chepstow in the Bristol Channel. In many regions. heteropods and pteropods.4 m. Bottom animals. In the Mediterranean it is only 35 cm. Where funnelshaped bays confine the tide wave. while are wholly wanting in the pelagial and others are represented only by a few aberrant forms. could not enter the abyssal depths until the pelagic animals and plants had established a food supply for them. and differ from them regularly by their swimming organs and arrangements to secure suspension. only highly specialized forms of worms occur in the open water.. at Granville on the west coast of 15. on the average.. 2 ' 3 The littoral benthos The littoral bottom fauna is the common mother of marine animals.4 m. up to The upper limit of the quiet water is also variable with local conditions. as in echinoderms and mollusks. Pelagic animals may always be traced to an origin from bottom-dwell- more than conditions. On the west coast of Europe the difference between tide marks amounts to about 2 m. divided into three vertical zones according The first is that of the tides. The depth of these zones varies with locality. a division. and in the Strait of Magellan. the tidal differences are much greater. supratidal spray zone may be distinguished above high tide mark. Such a region is spoken of as adtidal. to The littoral is motion of the water.

is coast from rocky the bottom is Littoral shores with loose substratum. the depositing shores Coastal areas with sandy or muddy bottom are grown over below the low tide mark." i. Movable. The distinction of loose bottom confined growth thus equally justified with reference to plants.. and the plant to algae and kelp. plants of . establish their roots and so take up the salts required by them. more or less loose beach material and solid rocky coast differ in many important respects in their influence rich plant life appears in coastal on ani- mal Below the low life.Littoral Shores with species Loose Substratum 221 and individuals in the littoral. with "seaweed. natural division appears to be that based on the nature of the substratum. were taken. like the animal Where the bottom is argillaceous or sandy. and at about 180 m. either with its environmental conditions or to the animal life dependon ent them. tide mark a marine strongly influenced by the substratum. with their extreme quent opportunities for isolation.. Marine mollusks are the most richly developed in the Indo-Pacific region. and their numbers are much greater than those of the pelagic forms. in about 900 m. those of the Philippines are estimated at 6000 species. great differences according to depth as the The is striking differentiation into favored by the great local numbers of specimens. Where waters. holothurians and pycnogonids for example. produces great differences in lighting. the great majority of benthic ani- much mals are found within this narrow coastal strip. with 33 species each in the Indian and Malayan provinces. Thus the littoral selachians are nowhere so richly differentiated as in the island archipelagoes of the Oriental region. line. in sheltered places. The dredge nets of the Chal- lenger rarely produced more than 10 to 15 individuals of a species from depths over 1800 m.e. on the continental coast vast numbers of single 58 The lists of species show almost equally as species were dredged. and the various combinations of the reference to The only conditioning factors produce a surprising multiformity in the whole. is extremely difficult. flowering plants can life. temperature.. The West Indies also have a great number of species. numerous species in the littoral fauna variation in conditions and by the freIsland regions. Although only 1% of the total ocean surface lies within the 180-m. afford the richest development of the littoral fauna. and oxygen supply. This is is hard such root functions are impossible. hundreds of specimens of single species. Its extension in depth to about 200 m. elaboration of coast line. motion of the water. A comprehensive characterization of the coastal area.

A very rich animal population develops in these thickets of seaweed. With so many herbivores. depending in general on the entrance of sufficient amounts of light. These are represented by various genera. appear in the Posidonia beds in the Mediterranean. and copepods. such as Ciona and Botryllus. which stirs up and redeposits the sand. A few lamellibranchs join this community. the related sea find. Blennius. feeding directly on the seaweed debris. Crustaceans amphipods. as do small sea anemones and the curious sessile medusa Lucernaria. support. which feed prithe on algal growth and slime that cover the living and dead marily leaves of the weed. the predators naturally are attracted in numbers. Gobius. Here also are to be found the polyps of the sessile generation of the medusae Aurelia and Cyanea. Numerous fishes depend on this fauna for their food. horses appear. swim among the * plants. in the same situations. the eelgrass Zostera in the North Atlantic. and are difficult to In the Mediterranean. and abundant oxygen. but less frequently from its living tissue than from the organisms borne on its surfaces and the detritus originating from it. Various types of worms maintain themselves on and in the bottom among the seaweed roots. this Common forms in biotope are the small snails of the genera and Columbella. The larger carnivorous snails. docea in the Mediterranean. and the crab Carcinides takes a variety of prey. As has been mentioned for Zostera these meadows of seaweed extend to various depths. and both individual and colonial tunicates. Animals find in them hiding places. isopods. with other snails. Bittium. The seaweed supplies food for many forms. . which is distributed by the movements 16 of the water through the neighboring area. or creep upon the bottom. On this algal growth there is a microfauna of protozoans. Posidonia and Cj/rao- and these with the addition of Halodule Such plants are wanting where there is too much wave action. and macrurans live in great numbers in the seasweed beds. and Halophila in the Red Sea. . Constant components of this biocoenosis are the pipefishes (such as Nerophis and Stjngnathus) which resemble the eelgrass leaves in both coloration and form. nematodes. The delicate branches of the hydroid polyps attach themselves to leaves and stems. but also dig down among its roots. Lobsters and certain crabs dig themselves burrows Rissoa. such as Murex and Tritonium. 82 '* 70 The inhabitants of the seaweed masses not only attach themselves to the living and dead leaves and stems.222 The Benthal the family Potamogetonaceae. schizopods. and the common stickleback are regular members. The starfish Asterias feeds on various mollusks.

19). This material is ingested together with the admixed sand by many forms. the holothurian Synapta. Others secure food with less Other mollusks. in the sand. 19. plants are available as protection and hiding place for the eulittoral. so do the lugworm Arenicola. and various sea urchins. admixture of sand by creating a current of water that brings them the finely divided particles of debris floating in the sea water. as well as feeding refuge for juvenile fishes.Littoral Shores with Loose Substratum eel. with which they search the surface about them. left. efferent or anal siphon to the right. the hemichordate Balanoglossus. Breathing is a matter of special importance to these sand-dwelling . as does the conger The seaweed forms a breedground and ing place for many fishes and for squids. draw in the fine deposit of food material on the surface of the sand by means of their long intake siphon. The great majority of these sand-dwellers feed on the decomposing remains of the seaweeds and of other animals. such as Scrobicularia (Fig. Amphioxus and many mollusks feed in this manner. just as earthworms take in their organic food with the earth in which they live. die wander slowly about. Scrobicularia piperata. though some Fiu. Animals of very different organization have become adapted to this mode of life and have acquired certain common characteristics. for the most part remaining fixed in position. the afferent siphon to the After Meyer and Mobius. 223 and live in them. a surprising number of animals of loose bottom areas secure their positions by boring more or less deeply into Where no animal life in the sand.

from the right: g. lh. gills on both sides of the middle portion of its body. bears a series of small spines along its arms.224 creatures. After v. <?. papillae on the protruded pharynx. Uexkiill. gills. by movement of its spines. 20. ing motion forces the water along 21) has a series of much branched its surface. of specially developed tube feet. tlie breathing and feeding chimney is reinforced by a secretion. duces a current of water that reaches kept open by means it through a chimney-like tube. pro- FIG. After excretory openings. 21. Id. 20). head folds. The heart urchin Echinocardium cordatum in the sand. p. the tubular arrangement of the spines and one of the prehensile tube feet are shown. The Benthal The lamellibranchs and amphioxus receive their oxygen with the water current that carries in their food particles. ventral setae. Astropecten The whose starfish vibrat- Fic. The sea urchin Echinocardium (Fig. The lugworm (Fig. dorsal setae. Arenicola. v. through whose extensive surface the haemoglobin of the blood efficiently takes up the scanty oxygen supply of its sur- . Ash worth.

23) the two large antennae together form a long tube for the intake of the and in the water for breathing. such as a dead fish. which burrow in the sand. its enlarged cross section of the breathing tube formed other annelids. . so that the crab can bury itself sand without being cut off from its oxygen supply. In the same habitat occur various portant. are recognizable by the spiral heaps of excrement pushed out of them. In them the water intake is situated in front. decapod Albunea (Fig. The number of forms concealed in the sand is anemones build Many astonishingly great. Turbellarian and nemertine worms conceal themselves in the sand.Littoral Shores with Loose Substratum 225 FIG. After Carstang. The arrows indicate the afferent and efferent openings to the gill chambers. both larger and smaller. Albunea symnista from dras. rounclings. tubes in the sand in which they can withdraw completely. and the water is conducted to the gill cavity by variously This occurs in Calappa (Fig. enters at the posterior border of the cephalothorax arranged channels. Mattila. Burrowing crab Calappa granulata from the Mediterranean. Among the annelids. In the anomurous is discharged forward. Mathe from side. dorsal An burrows. 23. the eyes may be seen on each side of the arrow points. front view. developed in the Singular breathing arrangements are box crabs. Arenicola is imsea FIG. and the claws just below the arrows. where the crab projects out of the sand. 21 In most crabs the water and Ilia. often closely crowded. After Carstang. 22. from which they may be lured to the surface by means of bait. 22). among which may be named the primitive Polygordius and by the antennae is shown to the right.

which establish the connection with the surface. The ventral siphon is the intake. . smooth-surfaced. and have siphons formed by the union of the posterior edges of the mantle into more or less elongate tubes. 24. They uniformly thin-shelled. in connection with their sand-boring habits. flat. bringing in the food and oxygen in the water. occasionally united. latter The agree in having relatively poorly bristle bundles. Protodrilus. in depth).226 The Benthal and the numerous nearly developed sessile forms. A curious companion of the worms is Balanoglosstts (Fig. ophelids. etc. After Stiasny. the terebellids. 19 and 25). while the dorsal cares for the outgoing current with the excrement (Figs. without bristles. in a U-shaped tube open at both ends and puts out piles of excrement are like those of Arenicola. and were formerly united into the group Sedentaria. clymenids. Balanoglossns clavigerus in its tube in the sand (30-60 cm. The mollusks of this habitat have much in common. a worm-shaped relative of the chordates which lives FIG. 24). chloraemids. the height of the heaps of excrement is 3-6 cm. have a well-developed foot without a byssus gland. maldanids. These siphons are convergent adaptations that have developed in phyletically distinct groups of mollusks.

in mollusks before the Cretaceous. A few * echinoderms are sand dwellers. 25 Fie. The irregular forms Brissop- Also Donax. Enlarged one and one-half times. The razor clam Solen (Fig. and Scrobicularia. are sand-inhabiting forms. Psammobia. Fie. in which the animal The retires from its surface position at any disturbance of the water. 26. 25. Tapes. seen from the edge. Razor clam Cultellus pcttucidus ( Solen pellucidus) of European seas. deep. Cardium. Cytherea. from the side. 26. After Francois. Mactra. FIG. drawn its into habit of digging tubes as much as 50 cm.Littoral Shores with Loose Substratum 227 in adaptation to life in sand and mud. and withThe opening of a burrow to the rear. is notable for The lamellibranch FIG. has. Venus and Tellina among others. a shell strengthened by its rounded form and strong ribbing. Siphons appear to be unknown * genera. its burrow. pismo clam of California burrows deeply into sand otherwise devoid of large forms. Lingula anatina in sand. After Meyer and Mobius. 25). in contrast. in the sand. which bores shallowly in the sand and which can jump by means of its angular foot. .

the thalassinids. The snail Natica (Fig. 26) are sand dwellers in the eulittoral zone of the Indian and West Pacific The brachiopods oceans.228 sis The Benthal (Fig. After Delia Valle. coming out at night to feed. square and 30 cm. 27). . Many brittle stars of the genus Amphiura live buried in of the ancient genus Lingula (Fig. the horseshoe crabs bury themselves in the sand during the day. 20) regularly live in sand. Haploops tubicola (Fig. and. FIG. The starfish Astropecten eats small worms. The sea cucumbers of the genus Synapta are also sand diggers and and Echinocardium swallowers. Amphioxus must be included in this list of sand dwellers. deep at the mouth of the Elbe. among decapods. lying in the sand with only the oral opening exposed (Fig. below the surface of the sand and attacks mollusks. and mollusks. Gebia in the North Sea. 27. A great number of crabs and small crustaceans dig burrows in the sand for example. the carnivores pursue their prey in this habitat also. 28). the sand. Haploops tubicola. Dahl secured more than 800 individuals of an amphipod (Bathy- poreia). Callianassa in the Mediterranean. A number of predators search actively through the sand for their prey. As everywhere. They live in holes into which they can withdraw by means of their contractile stalks. 29) moves about 2 to 5 cm. the last three are represented along the New England coast. ^chinoderms. between tide marks. which they with their food ingest supply as does the sand dollar Echinarachnius. boring through their shells and sucking out their contents by means of its proboscis. and the shrimp Crangon in the Baltic. Where they occur. By excavation of a space 1 m. Enlarged five times.

After Schiemenz. and the mouth also often turned up. which is of the greatest importance in secur73 The air bladder ing their prey. enlarged. They stir up the sand with a few undulating motions. Among rays. holding its prey with the divisions of its foot while boring tropical Trygonidae bury themselves in sand. otherwise sharks through the shell. Uranoscopus deeply. and when of their of their this settles it hides the outlines of their bodies.. and rays for the most part lie on the surface. and thus lie in wait for their prey. Numerous brittle . eyes usually directed upward. The number of forms living on the surface of the sand is small in contrast to the hidden inhabitants of sand bottom.Littoral Shores with Loose Substratum 229 Some fishes also hide themselves by burying wholly or partly in the sand. Slightly fishes the eyes are only slightly movable. with their fields of vision overlapping only a little (10-30). Trachinus and themselves majority of the bottom. their spiny Among the squids. 28. bivalve in sand. is absent or reduced in the fishes of this habitat. 29. This binocular vision probably makes possible an estimation of distance. adjusting its coloration to that of the bottom material. Aniphioxus Branchiostoma lanceolatum burrowed into the sand. the Natica josephina attacking a FK. Whereas in free-swimming carnivorous Fi<. the eyes of the flatfishes and of other bottom dwellers are very mobile and their fields of view overlap to the extent of 35-80. light-colored ventral side. All these have a flattened. according to the species. the flatfishes bury great lie The upon adapting their coloration to that background by means power of color change. Sepia tends to keep itself slightly embedded in sand or mud. protected by coat and rendered indistinct by their coloration..

and in them the tube feet are rounded instead of ending in a sucking disk.230 stars The Benthal and some starfishes have this habit. Fundulus. of crabs. require a broad foot. and this is the typical habitat of hermit crabs. one of the most common genera to be found along the Atlantic coast.). (). a contrast with snails in rocky habitats. from on loose bottom. Belone. along the coast of the Carolinas. Fishes of varied origin are provided with a rostrum with which they can plow the sand in search of prey. it follows its advance in over the sand. A number of carnivorous snails. where only a small foot is required (Fig." are said to dig up the bottom to secure 6:s their food. 30). and Inachus. The I They are attacked from the air smaller fishes of this littoral formation lead a precarious life. which often occur in great numbers. consolid bottom at 45-ni. e. A number FIG. roots out the Ammodytcs and anand Pegasidae feed in nelids. Hyas. with narrow foot. such as Buccinum and Nassa. such as Portunus.. on account of the looseness of the bottom. (/. there. After Hornell. 30. live on the sand surface. creep over the surface of the sand. Pyrnla sp. flattened foot. Such snails. living trasted with Rostellaria curta. as the tide comes in. especially in quiet water. with transverse teeth on their "saws. by terns and gulls and from the deeper water by larger fishes. The sawfishes Pristis and Pristiophorus. with broad depth. commonly lives in shallow water.g. and the halfbeaks ( Hemiramphidae ) the same way. rooting in the newly covered . 11 with its elongate lower jaw extending beyond the upper.

but the community conditions are relatively stable in their 77 instability.Muddy sandy bottom for food. In such places. The ecological processes here are in approximate equilibrium. the fishes collected in aggregations near the old outlet and. and in the main also the detritus from the rich litby toral vegetation. and neither organisms nor physical forces are changing the environment with rapidity. . It Deposits 231 continues with the advancing tide and often enters pools that retain the water for some time without holding it until the next tide. Mast failed to find any Fundulus stranded in these sandy tide pools.. this biotope is quite azoic. strings of islands. the minnows apparently test their line of retreat from time to time and return to feed along the bottom with their fellows. In a long series of observations. and at river mouths. passed out of the pool over the damp sand to the retreating sea water. This region of the sandy depositing shore is a climax formation comparable with the beech and maple community on land. Gravel and pebble beach composing the bottom has an important is found only in strongly moving water. Of course. Muddy The sea deposits amounts of organic materials constantly brought into the the rivers. since the movement of the stones on each other destroys all Coarse sand is poorer in animal life than fine. lies on the average about 200 m. close to the water's edge. are only slightly heavier than sea water and sink large very slowly to the bottom. When he dammed the outlet to such a pool. where the water of the opposing tidal current. and at much less depth in the Mergellina Harbor at Naples. life. The "mud line. but mud mixed with sand is still more so. below the surface. Muddy bottom is rich in life. He found that they could climb a ridge 10 cm. so that they come to rest only where waves and currents are absent. in quiet bays. and so is the animal life to be found there. 51 high and travel over land for 3 m. The effect size of the particles on its inhabitants. these deposits do not come to rest until much greater depths are reached. The deposition of mud occurs at various depths according to local conIt accumulates in shallow waters on coasts protected by ditions. there may be deposits of sand that tend gradually to raise the shore line and so cause the whole complex to move further out to sea." which marks the approximate limit of deposition of muds from the land. when the tide begins to ebb. individually or in groups. being stirred up by any movement. Mud on account twice daily stagnates flats in the Bay of Naples occur at 20 to 40 m. In the open ocean.

and other worms. Fades in depositing shore communities Although a general characterization of the fauna of the sea bottom with loose material may be made. ostracods. motility less.* The mud dwellers are on the whole more delicate than the sand A inhabitants: more fragile. These differences are excel- The mud by comparison of two sipunculid worms.232 I The Benthal affords a rich food supply. and Mullus barbatus. and the proboscis longer and a more slender. thinner shelled. Spio fuliginosus. but According to J. among fishes Botryllus aurolineatus and Ciona (tunicates). its animal life is not uniform. Zeus faber. The process of decay uses up the oxygen supply. but on account of the want of oxygen in their habitat there are fewer of them. Where the supply of fresh water is too small. . the musculature weaker. and Lophius piscatorius. Mixture of mud with sand is especially favorable and produces an increase in the animal population. and nudibranchs and turn. Capitella capitata. The mud line near the edge of the continental shelf is called by Murray the great feeding ground of the ocean. Tapes aureus ( lamellibranchs ) Bulla striata and Doris verrucosa ( opisthobranchs ) Nebalia galatea and Brachynotus . and swarms of small crustaprotozoans. Vast numbers of minute forms. Sipunculus nudus from sand and Phascolosoma vulgare from the mud. Bornia corbuloides. * sexdentatus (crustaceans). Degeneration and loss of the eyes is common. the skin less is softer. nervous system well developed. develop at the expense of the organic food materials and create a new source of food in their Mollusks bury themselves in the mud. mackerel and tunny. and even the whales. Arenicola. homogeneous mud. mainly by the oxidation of the hydrogen sulphide produced. . is for the survivors. Wilhelmi. crabs wander over its surface. more weakly muscled. Capsa fragilis. and isopods. . to which cod and herring. Certain lamellibranchs and the tube worm Spio fuliginosus are the most persistent sulphide. descend. yellowish gray or white coloration is frequent among them. Asterias tenuispina (starfish). the population decreases with the decrease in oxygen and the increase in hydrogen and deep black. sand. 29 The mud dwellers have mode of life in the main like that of the sand-bottom animals. nematodes ceans. and Spirographis (annelids). In the lently illustrated mud dweller. without admixture of most part without microscopic life. 87 the index forms of the mud fades in the Bay of Naples are Plagiostomum girardi ( turbellarian ) Audouinia. amphipods.

Fades in Depositing Shore Communities 233 changes from locality to locality in a notable way. Within this tially biotope special fades are distinguishable.m. in the Ringkoebing Fjord. of sea bottom. C. inhabitants of this area include. especially the water. Macoma comThe tide. 1 adult and 2 young Mya arenaria. 1 Aricia armiger. which are repeated A Stuxberg 81 single or of distinguished certain sections by the predominance of a few species. 31. FIG.25 sq. After Petersen.3 m. as yet unknown. I. G. even on extensive areas where the nature of the substratum and plant growth is essenuniform. Arrangement of animals on 0. exposed at low at other points when the same conditions are certain regularity in this diversified life is recogexactly repeated. between these facies and the depth of a connection nizable. and among annelids 4 Arenicola. depth. munity. among bivalve mollusks. must also exist. Petersen " has described the comr> . 2 Nephthys sp. and 1 adult and 3 young Cardium edule. at 0. 5 Macoma baltica. J. Other governing factors.

(12) Terebellidcs stroemi. 10. 2. in the Kattegat. numbered for identification. 4. (6) Aporrhais pes pelecani. (8) Chaetoderma nitidulum.234 The Benthd Arrangement of animals on 0. 32. and with the number of individuals. 1. texturata. sea urchins: (17) Echinocardium cordatum. 5.25 sq. ophiurids: (14) Ampliinra filiformis. FIG. pennatulids: (18) Virgularia mirabilis. II. worms: (9) Glyccra sp. 1. y (13) a fragment of nemertine worm. . 1. (7) Turitella terebra. (3) Cyprina islandica. 2.. (16) O. of sea bottom. (4) Axinus flexuosus.. 1. Filiformis community.. 5.. After Petersen. juv. (15) Ophioglypha albida. Mollusks: (1) Abra nitida. 6. (2) Corbula gibba. 1. 1. 3. (11) Brada sp.. EchinocardiumThe individuals.m. 60. 3. (10) Nephthys sp. (5) Nucula tenuifi 1. are listed below. at a depth of 20-22 in.

875. of them. (5) L. (7) Pecten septemradiatus. 1. The index forms of the different fades belong to different groups of animals. ophiurids: (16) Ophioglypha alhida. lamellibranchs (Fig. (4) Leda pernula. 31-34 give a good representation of four Arrangement of animals on 0. 30. worms: (8) Aphrodite aculeata. 32 and 34). of sea bottom. 1. (20) Morea Invent. 31). (6) Lima loscombi. 1. crustaceans: (19) Haploops tnbicola (see Fig. The inhabitants are: mollusks: 1) Venus ovata. After Petersen.25 sq. 1. sea urchins and starfish (Figs. (9) Glycera sp. (3) Axinus flexuostis. 27). 1. (15) nemertines. 3. robusta. 2. 1. (21) Verruca stroemii (on Pecten). FIG. I (II) fragment of a maldanid. or annelids may fill this role.m. . Ill. (10) Eumenia crassa. (17) O. 3. 33. 1. 1. 1. in the Kattegat. minuta. 2. 2. 7. (12) Pectinoria auricoma. 1. (13) a terebellid. Figs. (2) Cardium fasciatum.. crustaceans (Fig. 33). 3. (14) Balanoglossus kupfferi.Fades in Depositing Shore Communities 235 position of eight types of the hottom biocoenosis in the ocean surrounding Denmark. sea urchins: (18) Strongylocentrotns drobachiensis. Haploops community at a depth of 27 m.

(7) Nucula tennis. (12) Eumenia crassa. (Fig. with 875 speciall others are . Some of these communities are rich in variety. 8. 2. 31. In others. at a depth of 186 m. 17.. (11) Pectinaria auricoma. up to 20 species. (4) Portlandia lucida. (10) FIG. while others are poor (Fig.236 The Benthal and they are not necessarily the most abundant animals of the community. 5. in the Skagerrak. countless. Brinsopsiscommunity. of sea bottom. sea urchins: (16) Brissopsis lyrifera. 33). 1.25 sq. for example. (2) Cardium minimum. 9. 179.m. 14. (5) Leda pernula. The inhabitants are: mollusks: (1) Abra nitida. In some. with 6 species). predominates. a number of species are approximately equal in abundance. (13) Myriochele heeri. 1. so that Amphiura filiformis mens to 64 of the other species combined (Fig. 34. annelids: (8) Alicia norvegica. ophiurids: (14) Amphiura elegans. 20. 1. IV. Sarsii Melinna cristata. 8. (9) Artacama proboscidea. (3) Axinus flexuosus. 32) and finally Haploops. minuta. one species Arrangement of animals on 0. After Petersen. (6) L. 9. The facies overshadowed. 1. 11. (15) Ophioglypha sarsii.

especially juvenile fishes. Animals of this area must be able to withstand the temporary withdrawal of water. as platessa European food fishes attain their greatest abundance in the example Gadus aeglcfinus and G. as urchins Echinocardium and Spatangus. tinental shelf.Fauna of the sea of the Tidal Zone 237 bottom bear some relation to the distribution of the fishes. Sipho.* 3S> Fauna of the tidal zone tide mark upward the littoral fauna becomes more and more impoverished (Fig. as shown by the application of methods to the study of the animal communities of the Sound benthos by Shelford 78 and his students. the do the irregular ophiuran Ophiura ciliaris may literally cover the surface. Where shells of mollusks are heaped up on the loose bottom. they are exposed to the heat of the sun in the summer and to extremes of cold Food supply is also much less abundant than in and in winter. sessile animals take up residence. 31). The tides uncover a belt of varying width twice daily. of a country Most so-called coastal fishes are caught above the conOne can gain an insight into the wealth of the fisheries by merely examining a bathymetric chart. affording rich food for fishes. starsea Astropecten irregularis) occur at all depths. small caprellids and nudibranchs. whose stomachs are often crammed with them. adding to the variety of animal life in this habitat. rains subject them to fresh water at times. These facies established by Petersen have a wider applica- Petersen's than to the Danish waters. North Sea. From low * A number and P. The fish fauna is rich in this region of shallow water. especially hydroid polyps. in general much more so than over deep water. and Buccinum ) are abundant and widely disfishes (Asterias rubens. Pleuronectes microcephalus. The bottom of the North Sea 59 coastal conditions are not confined to the is The life is predominantly loose. on account of less favorable conditions of life. tributed. though the smaller amphipods and Cumacea are abundant. Its characterized largely by the abundance of echinoderms. Certain snails ( Neptunea. tion immediate vicinity of the coast but extend into shallow water far from shore. with their associates. The higher Crustacea are few in kinds and in individuals. and Rhombus maximus. and clay. certain fishes being found principally in or over similar areas. although never as index forms and always free to move over other types of bottom. composed of sand. frequently to the edge of the continental shelf. of for . sandy ooze. merlangus. but further inPuget vestigations are required to determine their validity.

the is reduced by the existence of and In with local ecological factors. the numbers of these worms per square meter Light is may be as high as 3. including diverse annelids. and form dark green bands. Terebella. and Echiurus. similar Littorina.350. and the quite resistant Cardium edule. This is true of easily as the stress of competition restrictions. balanoid. and laminarian zones have been described for the Pacific coast of North America 68 and for South Africa. 80 and can be recognized elsewhere sometimes with but slight modification. intermediate. and the Channel Islands. as do many sorts of crustaceans. The spray zone extends still 1 ' 1 higher. Basing his analysis upon his esting own extended study of the South African intertidal zonation in comparison with "the useful and intervolume by Ricketts and Calvin entitled Between Pacific Tides" Stephenson makes a summarizing comparison: The parallels between the two coasts as far as animals occupying similar "ecoMany of the photographs logical niches" are concerned are amazingly close. A more number of animals maintain themselves in this situation. Zonation occurs correlated with the duration of exposure to the atmosphere in the short. particularly with the substratum. Additional zones may exist extending over long coastlines. and of such mollusks as Venus.238 The Benthal below the zone of plant growth. similar habitats but that forms belonging to different species and often to different genera occupy and have a similar appearance. required for the mutual istic The annelids Arenicola. physical and chemical The biota varies with climate burrowing sea anemones. especially . England. Thus seven zones of algae grow one above the other along the shores of northern California and Oregon between the lowest of tidal low water and the highest high tide mark. Shallow tide pools retain shrimps hermit crabs. only euryhalinc and eurythermal animals are able to live within this belt of emergence. illustrating the Pacific coast fauna could be labelled with the names of South African species occupying similar habitats. without anyone but a specialist being It is not that the same species occur on the two coasts far from itthe wiser. of various kinds of worms. often for many kilometers. cycles occurring during the year. By count. In consequence. and the hemichordate Balanoglossus inhabit the intertidal beach. Nereis. is possible burrowers which below in living burrowing many biotopes low tide also dig themselves in between tide marks. 40 algae-like flagellates contained in them.000. Mya. and various other crustaceans. or long tidal 10 From the upper tide levels down. Vast numbers of small green turbellarian worms Convoluta roscoffensis are exposed at low tide along the coasts of Normandy.

Where a sand beach is freely exposed to surf. amount when the wave motion The California pismo clam lives is in Life of the spray zone Marine life. the fauna is enriched by the sponge Suberites domuncula. composed of a mixture of fresh-water and marine animals. they are stenobathic and confined to shallow water. in this situation have adopted an air-breathing existence and must be protected from drying out by a strong shell. in the region of exceptional tides. where The marine it. the conditions for plant and animal life are unfavorable. in considerable reduced. and the snails Melampus and the abundant Littorina litorea. the shores on each side are steep as high as ordinary tides reach. i. in the northeastern Atlantic. the supratidal zone. Change and by the tides has a median On these slopes is deposited a sticky and often very which a special fauna exists. the annelid Nereis cultrifera. and the widely distributed hermit crab Coenobita. Where seaweed is present.e. only spray and waves reach extends into a zone above high tide mark. Small quick-jumping amphipods (Orchestia) live beneath seaweed thrown animals that live up on the beach. the soft-shelled clam Mya. nudibranchs. and the ( See also Chapter 17. This applies to the above- mentioned Convoluta. ) ever-present beach crab Carcinides maenas.* 9 mixing of fresh and salt water and periodic stirring up I produce special conditions. Animal life is then usually wanting in the upper (terraqueous) levels and only appears this habitat. by the crabs Grapsiis and Ocypoda. so that the sand is continually stirred up and redeposited and the detritus particles washed out. finally. and more sloping above mean high tide. A river mouth at low tide channel. it includes.Estuaries 239 by the tides are so A number of species to its of the area exposed adapted special conditions that they are typically found in it. The list of marine animals is exdense mud in tremely small. Estuaries Estuaries form a special biotope in the littoral zone. Some of the related forms even extend further inland.. at considerable depths. anemones attached . which are air-breathers. a few Gammaridae. like Talitrns locusta. the worm Arenicola. Small snails are attached to the seaweed In the tropics the supratidal stage is characterized above all leaves. and the spider and hermit crabs with sea to their shells.

by roots and a variety of plants with similar growth habit. are consequently essentially fattening establishments for oysters (oyster spat) the oysters. Periophthalmus. in little ~ and floated in by the found a remarkable as- pools near the mangrove roots. The small semblage actinian Thelaceros rhizophorae is found at the water's edge. beds brackish-water where a wealth of detritus is stirred up by the tides. in Java. tide. Mangrove associations special facies of the estuarine habitat is represented in the tropics flfl3a>:io>42 The mangrove. their nests of the tide. scavenging on the remains of in the mangrove above mark the extreme height lamellibranchs Euryhaline bury themselves in the mud. cultivated oysters develop a form characterized by quality and size of soft parts. They are placed on the dorsal side of the head. and Littorina form a series successively more independent of the sea water. fish called The so that vision above the water is possible when all the rest of the body . These beds. Oysters. de- At ebb marine life. ants abound on the mud. The holes scending to a of the fiddler crab Uca are everywhere. These oyster beds require constant replenishment by brood from salt water. Its eyes are protected against drying (in the absence of eyelids) 8 by a great development of the conjunctival sack. is coasts. and to shallow water. especially Sonneratia (Myrtaceae). entering brackish water or becomto the mangrove roots. 35). are Such muddy bays. adapt themselves well to mud. In and upon the mud is of marine. for snails Cerithium. Various types of tree oysters attach themselves y example. depth of 75 cm. France. ing terrestrial. until it down by rains decays. and Aegiceras ( A Myrsinaceae ) . widespread on mangrove the mudskipper. living more out of the water than in it. is world-wide on quiet tropical coasts. and terrestrial animals. the artificial culture of oysters in employed for on the Atlantic and England. otherwise inhabitants of solid bot- tom. Ostrea mytiloides. Avicennia (Verbenaceae). The Potamides. as in many amphibious animals (Fig. The tangle of roots holds debris brought tide. with a thin shell. Between and upon the roots creep uncounted millions of crabs. with its stilt-like the mangrove beach.240 The Benthal to the tidal currents. the fertility of the oysters in is insufficient to maintain them. fresh-water. usually adapted in coloration to that of the mangrove bark. With careful protection from their enemies. Hermit crabs live on bottom and roots. with its attendant microfauna. exposed coast of North America. brackish water. Psammobiidae and Solenidae from the sea and cyrenids from the fresh-water swamps.

two-thirds natural After Hickson. and a few mammals (monkeys) frequent the mangrove flats for food. A great number of birds. stones. 35. it hops about on the mud and may climb the inclined branches of mangrove. Altogether. . size. however. and in South America. where single shells of mollusks. rock outcrops. and a few adaptable radical animals. especially predaceous starfishes and crabs. Littoral with solid substratum: eroding shores of solid bottom. Periophthalmus koelreuteri. the difference and obvious. appears immediately community submarine rocky bosses that project from the ooze of the quiet water in the The fauna and breakwaters are juxtaposed A Bay that of their surroundings. of Naples support a wholly different population from There are a few forms common to the two is habitats. especially of rock. but have their residence in the adjoining forest. Its food includes small insects as well as crabs. Supported on its tail and paired fins. Pic.Littoral with Solid Substratum: is Eroding Shores 241 submerged. stands in sharp conThis is especially obvious trast with that of the loose substratum. or harbor works to the loose bottom.. the mudskipper. quite different on a animals such The of foothold. certain reptiles. like Mytilus. the Indian and Pacific oceans. The mangrove in strand has a very similar development of animal life west Africa.

Bryozoa are predominantly rock dwellers. Tunicates often occur in gaudy carpet thick masses. Actinians. or the budding. also are more numerous on rock. especially those with calcareous tubes. Many tubicolous worms.* since in the quiet water of the deeps all available solid support is covered with ooze. relatively few animals have been able to acquire this habit. and these inhabit ma . Many of these sessile animals have the power of reproducing by shells of mollusks. Of the mollusks. close to the surface in the North Atlantic and Mediterranean the leathery tubes of Sabellaria cover low rocks so thickly that the openings of the tubes are juxtaposed. common on the rock habitat. and lamellibranchs in muddy sand. Those that live on sand are adapted to it by changes in their ambulacral feet. in general. but their tion. similarly Serpula tubes may form a tangled web among which the gill-funnels appear like colored flowers. Echinoderms. Broad areas of a rock may thus be covered by a single species of animal. and the The green and red surface appears like that of a honeycomb. Hydroid polyps form miniature forests on stones and rocks. The foot may be re- duced in the rock-dwelling forms. sand is not favorable for them. never in the sand dwellers. forming colonies of numerous individuals. 'The broad foot of snails functions on solid material. so common among the sand inhabitants. Lamellibranchs that live on rock usually have no siphons. majority scarcely requiring other protection against predators than their spicules or stinging cells and their powers of reproduction and regenera- one leaves the coast and as the water deepens. with their ambulacral feet probetter vided with sucking disks. snails and chitons predominate on rocks. Since this requires an enormous amount of work. require a solid basis. Hydroid polyps disappear 8 as small size enables them to gain a foothold on the leaves of plants. The of sponges and of sessile coelenterates are confined to rocks. are. with their creeping foot. carapaces of crustaceans. Tubicolous worms may cover wide areas. The great majority of simple and compound tunicates inhabit rock bottom. nly limestone or tufa.242 Sessile The Bcnthal forms are especially characteristic of the solid bottom. whereas their foot is provided with a byssal gland whose hardened secretion forms a fibrous attachment. sea anemones Actinia equina make a on rocks of the Helgoland. Never- . Some animals living on rock bottom have adopted the mode of protection furnished by boring into the substratum. the beautiful orange coral Astroides covers the rocks just below low tide mark in the Mediterranean. or similar softer rock.

Shells shells are thick-walled. Other rock inhabitants obtain protection through protective resemblance. thus Petricola. The spicules of sponges and the stingSnail ing cells of the coelenterates have already been mentioned. Many The fishes are able to that of the bottom. characters may be a response to the physical rather than to the biotic environment. boring annelid worms from both 25 the boring lamellibranchs are well Chaetopoda and Sipunculoidea. and the power of color change protects it when adapt their coloration to scorpaenids. and with their dermal appenpress wonderful color resemblance. of lamellibranchs are thick. their to close themselves support. and often covered with folds or spines. and the sea urchins are enabled to live between tide marks. such rock-boring forms are found in a large number of animal There are boring sponges. The holes provided by these rock-borers. like all other crevices. the size of which is adjusted to the growth of the animal. strengthened by corrugations. Pholas. which maintain themselves among . are made use of by other forms. appear their dages of varied form and it changes location. thus Echinus militaris9 Strongylocentrotus lividus (which does not bore in the Mediterranean!). and Heterocentrotus. the peculiar echiurid worm Bonellia. These shell this group. Cidaris. arched. Veneru- Eastonia. The animals living unconcealed on rock surfaces are often provided with means of defense. tough. The ark shell. where at the surface they would be unable to withstand drying. These borings are rock when exposed to wave action. known and are derived from various families. for example. Saxicava. Lithotrya bores in rock.Littoral with Solid Substratum: theless. Crustaceans have a thick. the "shipworm" Teredo. like pieces of alga-covered rock (Fig. and often beset with spines. lying in wait for their prey. and Alcippe into small shells. Gastrochaena. 33 be separated only by thin walls. and Lithodomus. spiny The name "echinoderm" indicates the nature of protection in shell. Barbatia barbata. Among barnacles. and Maia. so that often so as to numerous they are confined to this location for life. Eroding Shores 243 groups. Sea urchins bore even into lava and metamorphic pis. the Octopus sits in openings among the stones whose color is^ imitated by its skin. Inachus. and species of Arbacia. Hyas. 36). Such intertidal borings are founch only so placed that they retain water at ebb tide. the external openings of the tubes are frequently narrower than the tube itself. and the small claw-snapping decapod Alpheus are especially common in the borings of Pholas. Many crustaceans. at least in the sea urchins and starfishes.

On the coast of Chile Macrocystis may reach the surface from a depth of 80 m. FIG. calcareous and siliceous . extending to a depth of 30-40 m. as they must also do when they moult. 12 At other places. this growth is limited to algae. As there is no op' ' portunity for roots to penetrate. brown and red and bright-colored calcareous algae may form forests its These algal grass did in a mixed growth. they will be clothed with it. even more than the eelprime. If one removes this covering. and the growth may be com- pared to that of a forest. The upper levels. Each will consequently adapt itself to its special surroundings. they will promptly replace it. occasionally down to 10 m. while if a varied algal growth is Crabs of the genus present. being protected by the stinging cells of their messmates. 36.. their carapaces will reflect its variety. plant growth begins near the low tide level in rocky areas wherever the surf is not too violent.244 The Benthal rocks covered with other animals. sometimes completely covering itself. are grown over with algae like Sargassum and Fucus. These may be extraordinarily abundant and large. which are held fast by its tube feet. abound with animal life. one-third natural neapolitanum. they are thought to be protected by its inedibility. Below these come the sometimes gigantic Laminaria. thus where the sea lettuce Ulva predominates. Toxopneustes brevispinosus bears on its upper surface a number of shells. After "Aquarium Dromia bear a sponge that exactly covers them. Dragonhead Scorpaena porcus. 15 44 79 As on sand bottom. Some sea urchins mask themselves similarly. as the stands of eelgrass compare with meadows. Hydroid polyps. Hermit crabs living in snail shells to which sea anemones are attached belong in this category. have the extraordinary habit of 4 attaching algae." size. and tunicates to their carapaces. polyps. sponges..

Littoral with Solid Substratum: Eroding Shores 245 sponges. Kiikenthal of life from the West Indian waters near St. and this is a normal habitat of oysters. and numerous crustaceans and pycnogonids. nemertine. This results in restriction of the fauna. and tunicates grow on the leaflike blades. A number of urchins. especially below the laminarian zone. Malo. but remain alive. where they are reached by the high tide. their similarity of form. Stretches relatively close together may exhibit considerable Thus each of the coralline areas (Secche) of the rocky of Naples has its own peculiarities. and bathed in sea water for a few hours only twice a month. and free-swimming forms. In localities like St. and especially between tide marks. and its accompanying freemoving forms. which is greater the dinn 72 The more open the coast and the stronger the tides. protecting themselves against complete desiccation by the water retained within snails of the their shells. where the tides are especially high. small fishes feed here. Thomas: "At times our great trawl was filled with hundred weight catches of tunicates. force of the surf is greater on rocky coasts than on sand beaches. torn loose reaching the uppermost high tide mark. acles. is . As on the sandy bottom. small starfishes and sea chaetopod worms. At a slightly lower level. Sessile animal life. they may be attached to the rocks many meters above the low tide mark. sponges. They are exposed to the summer sun and the cold of winter. the These differences are separation. produced by convergent adaptation to similar conditions. Although these snails belong to different suborders of the gastropods. growth of algae is sparing or absent. tubicolous worms. and that the somewhat deeper-lying Scogli coralliferi in the same bosses on the bottom of the 48 Bay area harbor a different association of animals. They are mainly animals with a solid shell. All animals exposed to the surf are protected against being and crushed against the rock by special adaptations. as do small lamellibranchs with a byssal attachment. is especially rich in deep sounds where a strong current 45 reports such a wealth keeps the bottom free from mud. genus Littorina are interspersed among the barnacles. Lo Bianco says that the Secca di Chiaja is rich in animals elsewhere rare. All occur there. Nerita holds a similar position in the Indian Ocean. Schaugorgonids. Bryozoa. the fauna of rock bottom is unevenly dissorts of small creatures snails tributed. In the surf-beaten belt. differences. and and nudibranchs. turbellarian." more marked with greater geographic reports similar conditions in the straits east of Spitzbergen. especially mollusks and barnBarnacles of the genus Balamis extend to the highest level. hydroids.

The adhesion of Patella is supplemented by a glandular 54 secretion and resists a pull of 3. shell. Natural size. thickness. In addition. After Russell. 2 "' 38 This remarkable. 37). manent less its algal food. These mussels thrive best where FIG. to which it returns after short 69 excursions in search of FIG. and outlines of the rough and smooth surfaces. from they are most exposed to sheltered cliffs wave action.7 kg. because is Mytilus also thrives in places where the movement of water relativelv . where the surface is uneven the shells of Patella acquire a at their edges retain its corresponding irregularity (Fig. cm. per sq. 39 ). At the Fucus level individuals attach themselves to the rocks with the byssus. 38. with which they attach themselves to the rocks. for the most part they do not reach the laminarian zone. from the sole of Chiton pierced lead pipes 7 mm. Euand foot. their shells are pressed tightly to their supporting surface. in The form of the shell is also influenced of the edible mussel by the strength of the surf. chitons (Chiton). 38). Murex.246 The Benthal Below these appear limpets (Patella) and astonishing (Fig. upper. level. doubt- ropean seas. tending to be scarce or absent on is and islands (Fig. 37. Shell of Patella vulgata from the side. lower. so that a given snail must exact station. and Trochus at a still lower All these mollusks have a broad foot surface. the Indian Ocean. At such a perstation the rock surface is smoothed. with Haliotis. Littorina rudis. by the corrosive action of the secretion of the Instances are known in which the secretion Nerita funiculata.

Where wave action is weaker. however. In surf the individuals are small but thick- Wm Mytilus cdulis L. such as the starfish Asterias and the codling. at transitional areas thin patches of Fucus are interspersed with Mytilus colonies. algal growths of Fucus gain the ascendancy over them. fttt Cardium dut L Distribution of some index forms in the littoral of the Island of Bate and the neighboring coast of Brittany. Mytilus has disappeared entirely with the increase of the octopus/ 18 The surf zone. It is an extremely adaptable animal. so that the mussel in check. Main. 39. keep them shelled in the Bay of St. After Joubin. are closed to these predators. At other places enemies. and also the brackish water. as in Eroding Shores 247 the brackish water at river mouths and in some bays. FIG. B Haliotis tuberculata 4++ Tapes decussata L.Littoral with Solid Substratum: slight. or> . and with a strong byssus. and one may accordingly conclude that it is not the physical conditions but the biocoenotic relations that condition its abundance.

Its northwest coast is Haliotis in covered with barnacles and mussels. It is not unlikely that exclusion of enemies and more favored competitors by wave action has operated to preserve a number of primitive forms. 39 ) illustrates these conditions very satisfactorily. eliminate the competition of other forms. Chiton. and hence favors numerous animals that require moving water. and any protected sites on wave-beaten rocks afford a foothold to an assemblage of sessile animals and their free-living companions. obtusata."'' Hollows. and Haliotis. The outer face has only a few forms.248 The Benthal can flourish there. m. are primitive in structure and confined to the surf zone. supratidal stage. which reduce the numbers of species in unfavor- able sites to a few. foaming and oxygenated but without destructive force. litorea living near the low tide mark lays eggs that hatch into trochophore larvae and develop in turn into a veliger stage. In calm weather this water is warmed by the sun and concentrated by evaporation.000 per sq. m. neritoides. with specialized inhabitants. finally. The opposite side is washed by the water as it streams backward. cracks. L. living between the tide marks. supports a flourishing animal life of different forms. is present also on The spray forms small pools in hollows in the rock above high tide mark. The adaptation of Littorina to unfavorable conditions extends to its reproduction. Severe conditions. The selective action of the surf shows itself in other ways. while the inner abounds with animal life. and the common mussel. as these snails have become viviparous. An isolated wave-beaten rock affords different habitat conditions on its various sides. rudis and L. The small island of Batz near Roscoff ( Brittany ) ( Fig. The face toward the open sea receives the full shock of the waves. A rock coasts. in L. its south coast. these mussels form a connected strip half a meter wide exposed at low tide. Fissurella. the development is shortened. which are supplemented by below low tide mark. Nerita. in quiet water very rich oxygen. The same is true of groups of rocks and of larger islands similarly situated. Petersen 3580 larger and 95 smaller mussels per sq. 88 These progressive changes are marked by decreased fecundity. and the snails Patella. on long stretches of the Scandinavian coast. Mobius estimated the number of barnacles (H counted (Balanus crenatus) on a buoy at 10. in L. which live at high tide mark. the larval stages are done away with. the barnacle Chthamalus stellatus perhaps. while contftiued rains at other times . so that those few flourish greatly. the young leave the eggs in the veliger stage..

A few rotifers. annelids. Besides Littorina. affording a further example of the arrangement into communities described by Germany boundary. below low tide. contrasts with this type in occurring primarily on rocks. but active movement of the water is necessary to keep the detritus on which they feed in suspension. 71 Where the sand or mud placed on the shells of their predecessors. and crustaceans. and midge larvae Chironomus frauenfeldi are found in them. and are accompanied by a varied fauna of other mollusks. if stones or mollusk shells lie on the surface of more solid Australian Ostrea glomerata even attaches itself to living snails (Potamides ebeninus). It is sufficient for them. being living oysters. between Helgoland and the Netherlands- on coherent sand. 4 or 5 individuals often fasten themselves to the same snail. oysters do flourish. and sr> 05 and others. and forming reefs composed of solid masses of oysters that may be more than half a meter in thickness. The conditions in such pools are the more unfavorable. they cover a a depth of 34 to 42 m. not since it deposits on them is subject to motion. Ostrea glomerata. the higher above tide mark they occur. are also situated lie at breadth of 15-20 km. are distributed uniformly over the rock. their sand or mud. the uppermost 20-30 cm. on the Holstein banks are not close set. Oysters require solid bottom on which to attach their shells. a few copepods and isopods. There the pearl oysters Pinctada vulgaris. small and active isopods of the genus Ligia occur on the spray-covered rocks above the water line. pearl oyster banks of tropical seas are also associated with On the coast of Ceylon 28 such banks are found at depths of 10 to 20 m. The oyster banks of the deeper waters in the North Sea. sea anemones. a few snails. Petersen. The weight becomes too great. the snail is forced into the substratum. sensitive to reduction of salinity. Only especially euryhaline and eurythermal animals are able to survive in these pools. With the growth of the oysters. Oysters and pearl oysters. as support is available..Littoral with Solid Substratum: Eroding Shores 249 reduce its salinity. in Queensland waters. which are of commercial importance. The rock bottom. 56 hydroids. attached by byssal threads. They are not Verrill. are generally more or less abundant on rocky coasts. at places where sand and mollusk shells and organic materials have been consolidated by carbonate of lime. to be sure. aided by Bryozoa and calcareous algae. and buries them.. Oysters live on such supports on the banks of the shallow Holstein sea only 2 m. and dies. but spaced The oysters about a meter apart. where they occur on bosses of calcareous algae or fragments .

Coral reefs if a storm stirs up the sand Coral reefs form a special biotope in the rock-coast formation. Distribution of pearl oysters on rock bottom (left) and on sand bottom After Herdmun. below). Fig. important. Fie. 40. The Madreporaria (reef corals proper) are the main component of the coral reefs. coral may compete. (right). Coeloria.250 of coral. sandy areas may be fatal to the pearl banks and redeposits it on them. etc. the lime skeletons deposited by their ectoderm form the foundation. Fig. A rich fauna lives among the pearl oysters. anollusk-eating starfishes such as Astropecten hemprichii and Pentaceros lincki feed on them. extending from sea bottom to a little above low tide level. 41. sponges and Alcyonaria are abundant. The organ corals among Hexacorallae (Tubiporidae. and Nearness of annelids. known must be added to these. These reefs are raised structures. Three families of reef corals are particularly pora. the Acroporidae (Madrcand the Astraeidae (Gonastraea. to which the reef-building corals are confined. Fig. Red . with them for room. bryozoans. 41c) So must the Milleporidae. 41fo). formed by living animals and consisting of their calcareous skeletons. belonging to anto the inhabitants of the other class of the coelenterates.. they are assembled in clusters and modify each other's growth (Fig. Fig. 40). Great masses of limestone are built up under the favorable temperature conditions of the tropical seas. and tunicates cover their shells. The Benthol on the neighboring sand flats. the Poritidae (Forties. 41a).

Coral Reefs 251 111! .

eighty species of Madreporaria were found at Port Galero (Mindoro). which are more resistant. which is almost confined to the tropics. Calcareous Bryozoa compete with them in places in growing over and cementing The calcareous tubes of the fragile or broken coral branches.4 C.252 The Benthal Sea area as "fire corals" on account of their stinging powers. The solitary species Flabellum and Caryophyllia and the colonial Astroides occur in European waters. as in worms similar part. 30 A narrower equatorial belt. The variOne hundred and ety of species is also greater in warmer water. Foraminifera. but they flourish on the west coast. is the number of latitude. it is quite true. warmth-limited animals. pods. combined with the upwelling of cold water along west Africa and the west coast of South America. and in the Bermuda reefs (latitude 32 N. and 51 24 410 49 On the Tortugas. map. harbors all the large branched forms. in higher latitudes. (cf. The Gulf Stream enables them to reach and form reefs at Bermuda in latitude 32 N.) only 10 species of madrepores and 2 of mille57 pores have been found. On the other hand many corals are little sensitive to increase of temperature above their optimum. in the direct sun. Reef corals are stenothermal. is composed of carbonate of lime up to 90%. play serpulid 83 lamellibranchs sessile both and gastroSome Bermuda. Fig. Even within the tropics. The their bulk calcareous algae are important in reef building everywhere. limit the occurrence of coral.5. when they are exposed at low tide. at 25 N. Within this tropical belt further subdivisions (with reference to the coral) may be made. and Astrangia extends along the North American . areas of cold currents. ' ' Stony corals exist. mollusks. have principally the encrusting forms. also supply binding material. a sometimes an important one. may withstand a temperature of 56 C. and coral reefs are confined to warm seas. in spite of their minute size. approximately between 30 N. cementing fragments and filling holes.4 and 20.. Qoral polyps. with temperatures between 23. species scarcely 30.. species are known from Zanzibar. with water above 23. an evolution convergent with that of the reef corals. The cold southward current on the east coast of Florida prevents their development.5 C. contribute to the accumulation of lime. These usually subsidiary organisms are sometimes dominant in reef formation. These form massive branches by calcification of their ectodermal skeleton. 128 species have been listed from the very warm Red Sea. latitude and 30 S. while the two bordering areas. 77). withdrawn into their cups. They flourish only in water above 20.

Reef-building corals extend to a depth of 40-60 prolifera m. Broad gaps in fringing reefs occur and the great gaps in the Australian barrier reef seem to be opposite the river mouths on the mainland. probable oxide and of the wastes from protein metabolism of the corals. on account of the sediment carried in by the rivers of the sur- . Corals also appear in deep water as banks of considerable extent. zooxanthellae. The zooxanthellae were formerly considered to be the principal source of nutrition for corals in the photic zone. In the zooplankton at night. covered with Lophohelia and other forms.. It now appears that corals are carnivorous and feed on Digestion is rapid. They are absent in the region of the mouth of the Amazon and are much reduced or absent in some areas in the Java Sea. as for example the "Coral Patch" in the Atlantic at a depth of 800 m. its harbors. 84 is The growth of reef corals in waters close to the surface intimately filled correlated with the penetration of light. their is the removal of carbon difunction zooxanthellae. 20tS1 88 B0 91 Incidentally the presence of the * * ' zooxanthellae produces the bright colors of the polyps. Their polyps are with symbiotic alga-like flagellates. either embedded in the body wall or free in the stomach. are opposite the valleys that descend from the upper Egyptian mountains. though without forming reefs. 41 One may conclude from this fact that the rainfall in this region was greater in times not far past. depth. which represent gaps in the coral. 22 The same Lophohelia and Amphelia ramea appear on the Scandinavian coast north to the Lofoten Islands and are especially well developed in the Trondheim fiord. rarely down to 74 m. or red. starved coral polyps eject usually empty.Coral Reefs 253 coast from Florida to Cape Cod. yellowish-red. In spite of the aridity of the African coast of the Red Sea. coelenteron is relatively strong light of the upper four meters of water.. where they form banks at 200 m. which may be yellow. and during the day the In fact. at its edges. their abundance in the opposite river mouths. Corals from deep water have colorless polyps. The sensitiveness of corals to river water is increased by the fact that they are adversely influenced by the mud and sand brought down by the floods. differing from reefs in that they do not 7 rise to the surface. as is shown by Red Sea. Reef corals are sensitive to any decrease in salinity from the normal. the zooxanthellae produce enough oxygen in photosynthesis to fill the oxygen needs of the C orals. though increase in this respect does not affect them. which drops off to 1000-1500 m.

Other estimates are lower. A crabs. produced by Coral reefs frequently reach slightly above water level. land and snails. seaward. about 20 mm. bays and inlets where the tides are reduced. for the more compact forms. may have an age of several hundred years. 30 The rate of growth of corals is probably greatest near the surface. especially at capes that extend somewhat and on barrier reefs and atolls. an average annual increase in diameter of 39 mm. It is not rare. 42 ) and Millepora are the dominant genera. as do birds. however. coral growth In deep is weak. bringing with him the animal and plant human culture. which is constantly formed by chemical and wave action and by the activity of the reef-boring animals. for spring flood tides or great storms to wash all traces of life from such low islands into the sea. Such motion also distributes food and oxygen. Finally man associates of invades the island. which pile up loose blocks on the lowtide platform. and a vegetation may develop that increases in richness and The cocoa palm thrives especially in this situavariety with time. such as are valueless after 20 years. 80 The Benthal Coral reefs are absent in the neighborhood of active volcanoes. and this must be neither too active nor too slow. as in certain of the Tuamotus in 1878 and 1903. so that may be laid bare at ebb tide.254 rounding large islands. Too much motion prevents the ciliate larvae from gaining a foothold and carries them away. reach the new territory. 70 Arrangement of the types of corals in the reef is not uniform. Great blocks of hard coral. The rate of growth of the Maldive reef is ap19 Charts of a reef may be wholly parently 27-29 m. multitude of terrestrial invertebrates. The best collecting and observing stations for the scientist are always on the sides of the reef. Moderate motion of the water prevents deposition of mud. A certain amount of movement of the water is also necessary to the corals. insects.. Seeds drift to this beach and find a foothold. At the upper rim in the surf -zone Porites ( Fig. hold the erosion materials of the platform. small coral fragments and sand. and the accumulation may rise 1 to 2 m. "These alone seem able to withstand the force of the waves at their outer border. at the channels leading into the lagoon. the upper surface of the reef islands projecting tion. Coral above hightide mark are produced by the action of These storms. in 1000 years. The diameter of a large madrepore on the wreck of a ship sunk 64 years before was 5 m. and they form with the nullipores a . on each side. Many kinds of coral are able to withstand a certain amount of exposure. above water level for stretches of varying extent. since ash deposits kill them. the Astraeae. hermit crabs.

Outlines of the habitat forms of Madrepora muricata. and the great group Coral reefs are distinguished as fringing atolls. schcrzcriana of the Red Sea varies similarly. and cervicornis (Fig. above MeandrinaJ* 1 The branched madrepores have a different habit of growth at different depths. prolifera. After Vaughan. 42. They are often only a few meters wide. as do some same Poritcs. barrier reefs. are not true "The Fic. it accordingly extends as a rule about 30 cm. often within the species. of Astraeas. the rounded walls of Maeandrina. lines. as for example in Madrepora muricata with its three forms palmata. Parties sp. and pillars of Forties. and the last being ring-like coral islands. Fringing reefs are juxtaposed to the coasts and follow their out- FIG. when their platforms are exposed. They may be reached dry-shod from the shore at low tide. c. Astraea may also grow in shallow water and may be exposed at low tide without injury. according to the slope of the coast on which they are formed. 43. M. cornerstones of the reef are supplied by the massive coralsthe balls After Vaughan.Coral Reefs 255 breakwater which prevents fragmentation by the waves" (Darwin). b. but may reach a breadth of a kilometer. forma cervicornis." 41 reefs. forma prolifera. 43). a. more and more loosely branched as the depth increases. The madrepores rock builders. the narrower reefs are on . forma palmata.

46. The island of Bora Bora. (2) that the growth is most rapid on the outer face of the reef.256 es Salaam. explained these varied reef formations by means of a comprehensive theory based on the following considerations: (1) that Darwin 13 --. in the Society Islands.. with its barrier reef. Tuamotu Islands. distances. with a perimeter of 121 miles. After Agassiz. Such reefs have a steep outer and gentle inner sages (Fig. 44). 44. and ( 3 variations in ) that the level of the sea bottom take place.^-. From the first consideration. 45."-!- "-"^-"34?-? The atoll of Pinaki. (Fig. 61 The Benthal Dar steep slopes. Atolls are ring.-liT""-^'. it contains over a hundred subsidiary islets and a lagoon up to 100 m.or horseshoe-shaped reefs. ^-^^ ~ ^ l^gilgl^^ FIG. is 42 by 32 miles.. slope. and often more or less interrupted by pas- from 30 to 50 m. the width of its lagoon varying from 38 to 150 km. as for example at varying width. But when . Sunadiva. deep that connects with the open sea by 40 channels.-: FIG.!^ '. or on the coast of existing land. 45). " . reefs cannot rise to the surface from the oceanic depths but must be confined to submarine banks. The largest atoll. reef corals flourish only to a depth of 40-60 m..~v:r ". Barrier reefs are separated from the coast by a canal or lagoon of Islands may be surrounded by a ring-like barrier reef Mainland coasts may be paralleled by them for great Barrier Reef of Australia extends for about 1900 km. its depth The Great ~>if:^'5!?. the reefs are consequently of the fringing type (Fig. extending only a few meters above sea level. After Agassiz. such as the tops of submerged volcanos.. 1). On stationary and on rising coasts. the broad reefs on gentle ones.

where the central volcanic part is reduced to is about 50 m. Exact studies of Pacific reefs. as from and in the Sunda Islands. browsing of certain fishes. barrier reefs.e. however. When the submergence is long-continued. - ' rtl effects of aerial and wave erosion. 46. Barrier Dar es Salaam to the Red Sea 23 reefs and atolls. outline below. 76 The Darwinian explanation. the volcanic peaks have disappeared. atoll. Semper 75 called attention the fact that atolls. II). a lagoon and barrier reef are formed (Fig. the island itself may disappear. fringing Cross sections above. remains (Fig. leaving only a low platform surrounded by a wreath of small islets. boring of annelids and mollusks. II. Origin of various forms of coral reefs according to Darwin: I. Careful observers who have dealt with this problem emphasize two hills. but may be formed where the shore line is stationary. and chemical solution of the calcium carbonate on the determination of the height and compactness of the reef to and on the formation of lagoons. west of Tahiti. are not confined to areas of submergence. 1. and a channel or lagoon without corals is consequently produced between the reef and the original coast.Coral Reefs 257 mainland coast or islands are sinking. and fringing reefs may occur in . however. appearance of the volcanic peaks may be seen. found wide Dana 10 believed that he could support it from his acceptance. 46. III). and only the reef. about 100 km. now an atoll. near Bora-bora. "Atolls are the grave stones of sunken islands. more rapidly growing edge of the reef is separated more and more from the shore. i. high. and it still has numerous proponents. the outer. III. investigations of many reefs. on account of its simplicity. in each case. the highest of which at Motu-iti. In the Society Islands the progressive dis- FIG. as on Tapamano. 2.. reef." All intergradations between barrier reefs surrounding islands and atolls may be found. 46. and that these have fringing reefs. It is true that barrier reefs and atolls are in general absent along rising coasts. Volcanic islands with surrounding reefs are found in the Gambier archipelago. harrier reef. sea level in I and II respectively. have shown that the situation is more complex.

ways in which the balance may be established between the constructive and destructive agencies has as a consequence a great variety of appearance in the reefs of a single region. 1 Similarly the broad fringing reefs of the Fijis may emerge into barrier reefs with narrow lagoons. but Davis mature review of the whole problem in 1928 decided that in general Darwin's theory still represents the known facts better than any other proposed to date. according to local conditions. 19 "Glacial control" . Thus the so-called glacial control theory 8 takes cognizance of the Pleistocene ice age during which much water was piled up as ice on the land. and is understandable only from its own special history. and secondary lagoons. solid or loose volcanic material.258 The Benthal the same region in relatively close juxtaposition. shallows. formation. which tend to fill the lagoons and produce extensions of the shore. especially in the Indo-Pacific. tertiary limestone or The abundance of concomitant factors and the various modern coralline limestone. the product of a whole series of factors. The variety of material of the shore. The original subsidence theory appears oversimplified and needs extension to cover the evidence that water levels have varied through other processes than simple submergence. in 1875 it was an island of 9 m. west. and is now reduced to a submarine platform 30 m. The great effects of marine erosion are shown by the Metisbank in the Tonga Islands. and breccias is a factor that affects reef building. islets. under the influence of the strong trades. and the ocean level was lowered sufficiently so that submarine platforms could be waveeroded.. whose origin through erosion may be plainly seen through its gradual transitions. Rapid uniform elevation. below sea level. and south sides are surrounded by a broad fringing reef. all of which the existence of a wave-cut platform upon which coral postulate certain regions. On Tahiti the north. a deep lagoon has been formed between reef and coast. Every coral reef is an individual similarly produce varied results. Such platforms have since been covered by water released from glaciers to about the depth actually supporting coral reefs in is one of the theories of planation. it was later raised to 40-50 m. conglomerates. The uniformity of the whole 14 in a process postulated by Darwin is not valid in detail. These eroding influences are counteracted on one side by the growth of coral and on the other by the accumulation of coral sand brought in by waves and dust blown by the wind. while on the east side. height above sea level. or interrupted or slow elevation.

crinoids. Griffin reports from the coral reefs of Port Galera. skeletons of plankton organisms. (d) from the growth of deepsea corals.Animals Associated with Coral Reefs have developed. but the Tertiary lie above 725 feet and possibly as high as the 425-foot contact may On the whole a period of elevation is suggested. and holothurians. . (c) from accumulation of. 70 Boring through coral reefs comes from evidence secured by borings. 74 Hence results from the Samoan borings not only show planation but can be used in support of Daly's theory of glacial control.). is . and 200 to 250 species of crustaceans. The simplified. have been identified from a depth of 930 feet. 259 in Also. and 10 sipunculids. Subsidence tested. few starfish and sea urchins. 260. among them 25 to 30 . though planation and elevation may also have been factors in supplying a suitable base on which coral reefs could grow. above sea level. or (e) from some combination of these possibilities. This platform lies within the depth to which the sea level might well have fallen during Pleistocene Contours of the littoral sea bottom in the East Indies glaciation. Animals associated with coral reefs The coral reef forms the background and basis for a wealth of animal life unequaled elsewhere even in tropical seas. since Some complications there m. but numerous brittle stars. that the lowering of the sea level there at that time was about suggest 100 in. 8 deny. 111 species of Alcyonaria. may well have occurred in each of the other reefs are furnished by the Bikini borings. 70 species of chaetopod worms. as others suggest and areas in which the sea Darwin did not bottom becomes Such elevation might result (a) from crustal movement. over-brief. tabular summary shows that the three borings on Samoa reveal a wavetest of these theories A cut terrace in the underlying basalt. reefs may form reefs 70 sufficiently elevated. 3 echiurids. summarized in the table on p. Mindoro. the data furnished by these borings give much support to the subsidence theory. level" (130 m. (&) from submarine volcanic eruption." 19 74 In fossils - summary. . and this possibility is supported by the fact that Plio-Pleistocene coral reefs exist on the island of Timor (East Indies) at an altitude of 1300 in. ) evidence that the interval from depths of 300-575 feet (91-175 "was above sea level for an appreciable time after deposition. ( = 328 feet). "There are numerous instances of late tertiary and quaternary coral reefs being raised to great heights.

with smaller cracks and interspaces. The surf produces and brings detritus and plankton.Pleisto- Borodina) Below 1296 cene ft. 124 ft. afford retreats and hiding places to associated animals. 124.260 The Benthal Depth Reached. producers. 190. 2556 then subsidence Lower Miocene hermit crabs. connected with each other and with the sea. ft. 1947 (5) 118. Mobius estimates the number of molluscan species (without nudibranchs) on the reefs of Mahebourg. brown. Below 1020 Tertiary Below 1790 then ele- vation. all these. 124. 506 ft. Mauritius. Subsidence Reef 87 Kita-DaitoZiraa (North 47 1934-1936 (1) 1416 Above 340 cene ft. Even . The conditions that permit of such a rich develop- ment. which serve as food for others. in feet (num- ber of wells Location Funafuti Atoll. 1407 Not stated Probably not reached Subsidence near 47 Maratoea Bikini Atoll. there are broad caves and well-like hollows. Marshal] lands *7 Is- 300. and are fed upon to some extent by the The colorful mixture of green. Samoa 8 Recent 68. and red algae supports another series of animals. Planation: 124 wave-cut basalt terrace Great Barrier 1926. In general character this fauna associates it with the fauna of the rock bottom. at 336. 1937 (2) 600. Recent Not reached ft. The reef corals with their symbiotic organisms are to be reckoned among the food reef inhabitants. Ellice Is- Date 189&-1898 in parentheses) Geological (1) Age Reef Base Indication 1114 Recent Not reached Subsidence or talus lands w 1920 (3) 68. Subsidence. The coral reef habitat offers much better protection and concealment than solid rock. Upper OligoBorneo Shelf Islet 1947 (2) 856. It is permeated with pores and cavities like a sponge. and predators follow the herbivores. Not reached Subsidence PI io. in addition to the generally favorable tropical environment. as everywhere. are the abundance of food and adequate protection. 732 Recent 378. 1346.

to life in the reefs. As if there were coral. for which teeth are not needed. A number of coral-boring forms are represented. Numerous gastropods are sessile. not enough natural hollows. 17 Besides the above-named Magilus. 207).2 m. which is remarkably transformed by its life in the coral. The distribution of the gastropods in the Indo-Pacific is governed by that of the coral worms the older portion of the shell with lime (Fig. are abundant on reefs and may play a on which the molluscan fauna is especially varied. surface 0. they form a special gorgonid zone. below 10 m. in the Tortugas. it changes its direction of growth and extends the shell more or less in a straight line. which reaches a weight of 250 kg. especially in the West Indian reefs. Sponges of all sizes . Much of the animal population of the surrounding area will be concentrated on and in it. even 1. are grown over by coral. in order to maintain its contact with the outer world. Among the gigantic Tridacna. by a remarkable mode of reproduction. which live concealed in the reefs. Poterion. others that merely inhabit the reef without contributing to its structure are present. up to the great Neptune's cup. As in general on solid bottom. such as Lithodomus. the different forms belonging to it having lost their radulae in adaptation to feeding on the coralline slime. shows how attractive it is to other animals. among them the gigantic discosomatids with an oral (Stoichactis) in diameter. the family Coralliophilidae is especially adapted reefs. pearl oysters. beginning as a snail with ordinary spiral shell. a whole series of forms bore into the The examination of a solitary madrepore. sessile animals are present. It must suffice to emphasize the most important common characteristics. and others. is an index species of the Indo-Pacific reefs. like Magilus. the lamellibranchs here characterized by special thickness of shell are oysters. as it becomes overgrown.Animals Associated with Coral Reefs 261 the individual coral heads are porous. located at a distance from the reef on sand bottom. The snails of the coral biotope are remarkable.. Serpulid part in their structure. filling 17. Gorgonids may be abundant. occur. are characterized Some predaceous chaetopod worms. In the palolo .. Sea anemones are numerous. 4r> There is scarcely a benthic group of animals not represented in the coral communities. p. and when one is broken up a crowd of varied inhabitants scatters in all directions. numerous In addition to the reef-building corals. like Coralliophaga. and the spiny-shelled Spondylus.5. such as the solitary proteoid Fungia. in cracks in the coral.

of the worm. and these parts are cast off when mature. when they leave the reef and swarm out into the open surface water. 47). variously concealed or in cracks and between the branches of the madrepores. Palolo worm Eunice reef. and Tr. long-tailed their distribution. decapods that make a snapping sound with their claws (Fig. Wood worth. for example. Crabs arc abundant. The alpheids. at a predictable date. FIG. pistol crabs. namely on the day before and the day of the last quarter of the moon in October and November. 49) confined to coral reefs. The larger species live in the deeper channels and swim in and out. as more slender posterior portion breaks off and swarms in the open water After as the palolo. the deep body form with . where single The thick anterior remains in the end and the species are restricted to single reefs and occur only on single kinds of corals. Many of these fishes have a coloration like that of coral rocks. 47. guberrima on Madrepora hairnet Trapezia on the Uyanga reef.262 The Benthal viridis (Fig. yellow and green sharply contrasted with their background/' 3r> Certain types of pattern and body form are repeated by genera of distinct families. Eunice South Seas the eggs and sperm are developed only in the posterior division of the body. is The family Trapeziidae (Fig. are closely correlated with corals in small. viridis. "The coral reefs of the South Seas literally swarm with fishes. the reef and remain in tide-pools at low tide. Out of 79 long-tailed decapods taken by Gardiner in the Maidive reefs. and remain in the protection of the coral. This occurs twice each year. 6 The number of species of fishes found in coral reefs is large. exhibiting The smaller species live on the surface of tihe most brilliant colors. Others exhibit so-called warning colorations. 76 were alpheids. scarlet. Eunice furcata has the same habit in the Tortugas/' Crustaceans are exceedingly abundant Dry 3 in the reefs. 43 The sea is then so filled with them that the natives gather them with baskets for a feast. especially the small Cyclometopa. rufomaculata on Pocillopora favosa at a reef south of Dar es Salaam. with blue. sometimes so strikingly that such fishes were formerly classified in the same group. 48).

In spite of stony skeleton and stinging cells. Crabs mask themselves with sponge. and the barnacle Pijrgoma. the lamellibranch Coralliophaga. Fig. and the suborder Plectognathi with the trigger fishes (Balistes. and the rounded or slightly concave tail fin. . 48. in rather large pieces.Animals Associated with Coral Reefs vertical black 263 bands and long median fins extended backwards. All the hiding places are utilized. as in many butterfly fishes (Chaetodon. Silver coloration. FIG. is extreme. Fig. 52&). spotted or vertically banded body with many striped head markings. 50a). boring lamellibranchs. more than elsewhere in the rock habitat. Competition. Among these are the pomacentrids (Fig. Trapezia rufofyunc- mudensis. natural size. 49. and the annelid Siptinculus). and Glyphidodon) Species of the percoid genus Serranus and some labrids may resemble each other in their elongate elliptic body form. and the puffers (Fig. is very rare among coral reef fishes. . 510) and pomacentrids (Dascyllus. After Miers. these fishes browse on the coral. Fig. Protective devices are accordingly evident. Fig. 50fc. the trunk fishes. 50c). Fie. 520. 50 Other fishes tion of the teeth into show convergent adaptation in consolidaa beak. intermedia. Pistol crab Alpheus hersize. of coral. Families in which a uniform coloration is usual contain fishes with distinct patterns in the East Indies. Holacanthus. enabling them to bite off small branches of coral. as in the banded Synaptura zebra and the spotted muraenids. has been found in the stomach of Diodon. uniform dorsal fin. Many forms perthe disused holes including mit themselves to be overgrown by coral (the snails Magiliis and Vermetus. About twice natural After Spence Bate. 50a. otherwise so widespread. tata var. the sparids (Fig. amongst so great a number of animals in a limited \habitat. up to 1 kg. long. Others bore into the coral sea urchins. of the deceased borers. Diodon and Tetrodon).

Trigger fishes lock themselves into crevices. which they resemble in FIG. b. whose emerald green be the case with the opisthobranch color. ally poisonous. a trigger fish. . which permit colors to vary relatively unchecked by natural of color selection. Batistes aculeatus. Reef fishes with similar patterns: a. a pomacentrid. many such fishes are actucoloration. The riot and pattern displayed by coral reef fishes may arise in part from the presence of rich food supplies and innumerable places of refuge. and Psenes) of sea anemones (Stoichactis and others). and Dascyllus aruanus. even seeking shelter in the enteric cavity. Tra71 seek protection among the stinging tentacles chichthys.264 The Benthal Fishes and cephalopods change color in correlation with the background. in addition to an unpleasant musk-like odor. Small shrimps and fishes (pomacentrids: Amphiprion. This may also snail Bulla. 50. The bright colors of some reef fishes may be interpreted as warning coloration. a butterfly fish. Cha&odon striatus. c. with orange spots. indicate its unpalatability.

a. and the gorgonians are poorly developed. a porcupine After Jordan and Evermann. a parrot fish. the poverty of the reef fauna is farthest east not everywhere the same. Tridacna of 15 to 45 cm. an angel fish. similarly impoverished. After Jordan and Evermann. Microspathodon with similar shape: a. and the accompanying fauna is The fleshy alcyonarians are notably absent. fish. few sponges occur. shares of species of corals is latter. total effect of a coral reef is overpowering . and fishes with teeth fused into a Leak: Sparisoma aurofrenatum. The toward the open Pacific. Holacanthus tricolor. than elsewhere in the same latitudes.Animals Associated with Coral Reefs 265 The richness of the Tuamotu Archipelago. are dwarfs in comparison with those of 1 Queensland. Gorgonians and alcyonarians are noticeably absent in Tahiti also. Reef fishes and b. a pomacentrid. 51. dorsalis. Diodon hystrix. 52. and the KJG. The The rich fauna of Indo-Pacific fishes reaches its maximum wealth in the East Indies and is much reduced in number of species in eastern in its Polynesia. The number smaller FIG. Reef b.

C. 1888. The Dutch flower/' refer to them in the East Indies as "gardens. C.The animal life of the lagoons of barrier reefs and atolls is considerably different from that of the outer face of the reef. "Studies in marine ecology: IV. The bottom mud Solitary corals are frequent in the lagoons." Museum Comp. yellow. W. The corals themselves. und Boleophthalmus) sarze: Augen des Schlammspringer (Periophthalmus Bemerkungen zu dem von W. "Studies in marine ecology: I. where every animal becomes a In spite of many common characters.266 of life The Benthal and splendor of color. 3. Zool Jahrb. text figs. 2.. of the reasons for the failure of corals to flourish in the lagoous. Fishes of the lagoon are mostly dull colored. red. of the lagoon is covered with coral sand and calcareous derived from the erosion on the reef. 1903. branched. "Die Maskirung der oxyrrynchen Dekapoden durch besondere Anpassungen ihres Korperbaues vcrmittelt. Volz verfassten AufZur Kenntnis des Auges von Periophthalmus und Boleophthalmus. While the reefs at Tur ( Red Sea ) marked by the predominance of warm colors. W. Agassiz.." Biol. 1923." limiting the geographic range of invertebrates of the 4. "The coral reefs of llie tropical Pacific. "tTber die 5. Alice. Aurivillius. 238 pis. millepores are the most abundant corals. in contrast with those of the reef." . . Anat. the general appearance of the reefs is not everywhere the same. 44:167-191. 23. L. Eine biologischmorphologische Studie. Handl. Baumeister. and are often slender. 35:341-353. They are less commonly continuous. Mem. The number of animals in the lagoon varies according to local conditions. and fragile. but form spots and accumulations in limited areas. 1913. Zool." Kgl Svenska Vetenskapsakad. Agassiz found the lagoons of Pinaki and Rangiroa rich in fishes. The effect of temperature in littoral. are malachite green Anthophyllae stand beside olive green millepores. No. 4:341-355. 1923. and brown. 4:1-69. BIBLIOGRAPHY 1. 1 fig. and this is probably one with Porites next. and emerald green madrepores and astraeas beside brownish green monti7 pores and maeandrinas.. Idem.. orange. 5 pis." Haeckel speaks of "fascinating coral groves.. where yellowish green Alcyonaria accompany sea-green heteropores. The distribution of common littoral invertebrates of the Woods Hole region. 28:xxxiii. have quite different habits of growth.. Bull. green is the dominant tone in the coral gardens of Ceylon. S. In the lagoons of the Marshall Islands. 410. in quiet water. Woods Hole Ecology. Alexander.

1880-1887.. B.M. 1908. with some remarks on the of a utility of specific characters in the genus Calappa." ct 17.. Zool. 408. Fischer. Charles. 239). 2 maps. London. (p. M. Rept. N. 1 fig. Congr. Y. and C. "Critical tide factors that are correlated with the vertical marine algae and other organisms along the Pacific Coast. 16. 15. sci. Jr. Arch. 3rd ed. Davis.Bibliography 6.. Gardiner. 18. 267 Beebe." Pub. "The building of atolls." Pub. 1910. map. 1940." Sci. 1912. wlss. 551.5-328. Coral Reef Comm." J. 3 pis.. 1931. illus.. 8 figs. Congr. 366. Anton. Dodd. with special reference to the Alcyonaria. 13a. L.S. 5 figs. 4 maps. London: 4th xiii. Paul. Natur- wissenschaften. . 15 pis. Sea. Hjulmar. 1905." Rev. Cambridge: 118-123. Sci. 13. 390).... 20. Monthly.. "Mitteilungen aus und iiber die zoologische Station von Neapel. 7 9. Idem. Griffin. Dnhrn. (4) 11. Broch. 27:31. Intern. 1901. Zool. Doty. 19. H. 1928. 31:14-30.. "The Philippines Marine Biological Station Port Galera. "The Roy. Beagle under the command of Captain Fitz Roy. "On some modifications of structure subservient to respiration in decapod Crustacea which burrow in sand. Cyril. Mindoro. Zool. J. M. Zool. and Coral Islands. Davis. 120). Gratz: 682-688.. f (5) 10:385-393 (p. Wash. 471). Cafy. "Studies on coral reefs of Tutuila. D. 1907. 1874.. Y..5-334. Gravier. pis. Dantan. L. 8. 10 ff. Appleton: xx.. Crossland. 23 pis. 10. zobl. "Riffcorallen im Nordmeer einst und jetzt. "Notes ichthyologiques. Murray: xvi. Walter. L. pi. exptl. 160 figs. W. Journal of researches into the natural history and geology of the countries visited during the voyage round the world of H. Our search for a wilderness: an account of two ornithological expeditions to Venezuela and to British Guiana. M.." 7. 27:289-300. Wash. Beebe. 9. American Samoa. Intern.. 1897. Corals Linnean Soc." de paleontologie ou Librairie Savy: Paris. 32 S." Proc." 3: Ixi-lxxvii. "The formation of coral reefs. Idem. Coral reefs and atolls. 1 map. 1-1369. 413:53-98. P.. . "Reports on the marine biology of the Sudanese Red Sea: IV. distribution of 1946." Z. E. R. 517:303-412.. "Sur quelques traits de la biologie des recifs coralliens. "The ecology and geologic role of mangroves in Florida. illus. (p.). 12 pis. Y. 1922. 23." Quart. 1-6. Soc.. Mead: 1-398. text figs. J. Idem. 22. I. The recent history of the coral reefs of the mid-west shores of the Red J.. Garstang. [Funafuti Reports] 1904. Carnegie Inst.. 21. 14 figs. Charles. atoll of Funafuti. 1913. J. pi.s. 12. 10:804-806. 40:211-232. Structure and distribution of coral reefs. Manuel de Conchyliologie histoire naturelle des mollusques vivants et fossiles.) new species of Albunea.. and the description (n. (p. 1898. 4 pis. 1895. 6:32. Dana. 14. 1138 figs. Rev... figs. S.. Hydrobiol. N. Ecology. London. 25:457-480 (p. 1. R. Darwin." at 1 Verhandl 8th 24.N. "Sur les recifs coralliens de la baie de Tadjourah (Golfe d'Aden). Microscop. 1884... N. Holt: xix. 181. et gen. Carnegie Inst. W. London: 1-428." Intern. 1931.

La baie de Saint Malo. 3 pis." Ibid. Kiikenthal. 221 figs. W. 33:379-420. 1912. 175 (p." 37. cotes rocheuscs. 26. 443-449). A. Forschungsreise in 1896. 115:1-20. S. John. 5 parts (pt. 1 fig. 27.. 1889. 28 111: 30. Jones. Lipsius & Fischer: xi." pi. 67:1-32. C. the Sanghir and Talaut 31.. A. Louis. 189). Proc. Krumbach. 36.. 1908... 25).268 25. Herubel. 35. 41fl. map. 392. "Etudes. Klunzinger. Willy. Karl. La Les repartition des animaux dans ses rapports avec la nature des rivages. (p. E. frontis. 44. 172:1-12. Report to the government of Ceylon on the pearl oyster fisheries of the Gulf of Manaar. 1903-1906. Berlin. Java zoologisch en biologisch. Idem. Cambridge. "Ueber bohrende Seeigel. illus." Bull inst.39. Nahrung felsenbewohnender Zool Anz. text figs. Jordan. Plant-animals: a study Cambridge Univ. La vie dans les oceans. 1889. Landbouw: 1-663 (pp. M. Idem. 15. France. Thilo. Kramer. 34. F. "Notes preliminaires sur les gisements de mollusques comestibles des Cotes de France.. 140-146). in the archipelago. ." 45. 5.. Paris. zoo'l.. D.. 726. Press: viii. 110. Issel. F. A naturalist in North Celebes. and Alvin found of the species fishes of Oceania. 33. J. 40. et la biostatique des sipunculides. Haeckel. Idem. 1924]. Ueber den Bau der Korallenriffe und die Planktonvertheilung an den samoanischen Kusten nebst vergleichenden Bemerkungen. Monaco. "Ergebnisse einer zoologischen . Zool Soc. map (p. Stuttgart. Ein Ausflug rwch Trieste und der Quarnero Beitra'ge zur Kenntnis der Thiertvelt dieses Gebietes. Augustin.. H. Levy & 400. 32. Buitenzorg. McGraw-Hill: xii. 25:173-455. 44:440-451. The Bcnthal 1861. Georg. J. Ill text figs. 1878. Monaco. illus. 415. Naturgeschichte. Flammarion: iii. "The fishes of Samoa: description with a provisional check -list of the Bull Bur. London. 1 text fig. 174. Idem. 1:268-302. Berlin. Koningsberger. xii. p. in symbiosis. N. (p." Ibid. della foglie di Posidonia studiati dal punto di vista bionomico. "Mitteilungen iiber die Seeigel der nordlichen Adria. II bentos animale Raffaele. 2 pis. J. 29. La cote nord du Finisterre. Bull inst. Herdman. Dept. 42. 38.. Gebruder Paetel: xvi. 334. M. 1 map. London. 35 figs. 1903. A. "Biologia neritica meditcrranea. 1909.. 1897. Indische Reisebriefe. illus." 1909:671-679. pt. Protozoa through Ctenophora. Koralltiere des Roten Meeres [fide Hesse. Hyman. Keeble. Bilder aus Obera'gypten. 43. 1940. 81). Nicolai: viii." 1906. Le Golfe du Calvados. Syst. 1906. oceanog.." Zool Jahrb. flora. 81). 1914.. comparees 124. Fisheries.. 71:1-24. The invertebrates. "The coral island question. oceanog*. Murray: xv. in Minahassa.. 445-448). 55.. 21 pis. 2 pis. 2 maps (pp.. a narrative of travels Islands. "Etudes sur les gisements de mollusques comestibles des cotes de France. L. S. Seale.. Bull soc. Hickson. 1912. 1906.. W. 1893. Kiel. Guerin. Premiere contribution a la ?norphologie et pJiysiologie 28. 1911-1915. 1919. Joubin. and ethnology of the districts visited. 1910. Miller: 41. 163. Arch. W. Y. "Considerations sur la faune des cotes de Franc* 1 .. illus..ip. 1 pi.. etc. with notices of fauna. Ernst. Grube. 22 text figs. 1879. 1 m..

Korallenriffe. Hamburg 2:117-134. 1875.. Marenzeller. Zool Neapel. Idem. "On coral reefs and other carbonate of lime formations in 61. 17:79-109. The depths of the ocean. 107:51-55. 13:157183. "Die neueren Forschungen iiber die Bildung der Zool Inst. E. 50. periodo di maturita sessuale degli animali del golfo di Napoli. Ges. Menke. R. 59. 521 ff. Idem. With contributions from A.. A." C.C. atoll. Moseley. L. H. modern 1897. and Robert Irvine. ihre physikalischen "Uber die Thiere der schleswig-holsteinischen Austernund biologischen Lebensv erhiiltnisse. 37:19-30. 102:105-112. F. Salvatore. 46. Carnegie 54." (Kiikenthal und Hartmeyer.. 1893:67-92. 1901. 3rd Intern. J. 1911. 1895. "Notizie biologische riguardante specialmente Mitt. 58. O.. 1892. 56. millan: xvi. 9:229-245. Karl.. H. Murray. S. 1908. 1902..N. R. Lo il Bianco. Syst. biinke. Thomson.B. "Ergebnisse einer zoologischen Forschungsreise 269 Abhandl Senck. "The behavior of Fundulus. 1907). and G." Ladd. illus. Mast." Sitzber.MS. Leyden: 99111. Edinburgh. Akacl Wiss. Beih. May. Gran." 48. 1892. Mielck. I. Tracy. 1 pi. G. with especial reference to overland escape from tidepools and locomotion on land. 57.N.. 55.. Jahrb. John.. 1948. Eduard von." 2 vols. Zoologischer Teil. von.. "The annual breeding swarm of the Atlantic palolo. 49. 51. Wash. Suppl. Dekapoden Familie Tra- peziidae. under the commands of Capt. S. 1899. Meeresunters (Kiel. Notes by a naturalist on the Challenger. "Ostafrikanische Steinkorallen. 13:448-573 (p. 5 pis. 620. Martens. "Die geographische Verbreitung der Ibid.S. 47. H. Ortmann. Appelof.. 1912. W. 53. Mobius. 5:341-350. (p. Helgoland). Centr. 119). Mayer. Marshall Islands. 62." Wiss. R. S.R. being an account of various observations made during the voyage of H." /. Lill. T. Zool..... Roy. illus. nach Westindien Zool.. "Die Hydroiden Untersuchungs fahrt des Rcichsforschungs dampfers Poseidon in das Barentsmeer im Juni und Juli 1913. seas. and Capt. K. F. Die Litoral fauna Ternates.Bibliography den Molukken und Borneo.). a general account of the modern science of oceanography. G. based largely on the scientific researches of the Norwegian steamer Michael Sars in the North Atlantic. 1 pi. "Drilling on Bikini Science. Abt. Ileinrich. in Scheuring. Soc." Sta. Murray. "Die Preussische Expedition nach Ost-Asien. London. 18. in the years 1872-1876. 11:1-12. 1893. 1922. Zool. Anst. Berlin. Helland." Zool Anz. . Congr. Nares. Murray and Hjort." figs.. 1915. 3 52. Einleitung und Reisebericht. "The general conditions of existence and distribution of marine' organisms." Pub. A." 10:201-216. 60. Jahrb. John. Mac- map. Challenger round the world. "Physikalische und physiologische Factoren bei der Anheftung von Schnecken der Brandungszonc. E. t 22:45-52." wiss. Jahrb. 1891." 6:631-670.." Proc. Animal Behavior. Appendix. "Die Korallenriffe von Dar-es-Salaam und Umgegend. 1909. and B. Sir G. Walther.

"Reisebericht.. 25:1-62." de la Polynesie. H. 5:1-122." Zool. Jensen. "The constitution of the intertidal fauna and flora of South Africa. "Evertebratfaunan i Sibiricns ishaf. 1913.. M." Linncan Soc. faune.. 77. Russell. grundade pa de zoologiska untersockningarna under Prof. "The sea-bottom and its production of fish food. "Some marine Coast of North America. London. Stuxberg. analyses. 14-15 (p. Carl. Warne: xi. seas. 67. illus. The Great Barrier Reef of Australia.. Ges. S.. J. Yonge. t 78. 16). pis. 9:235-247. E." Zool. mus. London. 21:1-44. "Zur Kenntnis der Azorenfauna. 320. .." Ibid. T. Natal Museum. 1943. 64 pis. 66. 76. zool. F. 1 map. 2:856-870. 'Siebold und Kolliker. H. 1899. 387.). 15-16. Pacific Palo Alto. Natur- geschichte. Forelopcncfa A. Danish Biol Sta. Petersen. "Vorliiufiger Bericht iibcr zoologische Untersuchungen in nordlichen Eismeer in Jahre 1898. 65:1-16. Ann. Arch. Ricketts." forsch. C. 1928. C. 16 text figs. and Dorothy 1942. 74.." Part III (Bibliography). The 127 pis. Idem. and potentialities. Pappenheim. G. Seurat. 35).. Monaco. structure. 1879. J. 1918. 112 figs. et 1935. F. pt. dcut. 64 pis. B... Anton." Zool. C. 3 charts. "Animal life of the sea-bottom. 65...270 63. 81. Great Barrier Tides.. 1863. "Zur biologischen Bedeutung der Siige bei den sogen. "Great Barrier Reef bores. Simroth. its products London. 1948. et Bull. 72. Schaudinn. tion of life in the [Indo-Australian] Archipelago. G. E. E. Pacific 608. London." 1926 and 1937: descriptions. 2 figs. und Pristis Lath. 1929. E. and P.). "Valuation of the Sea: II. 1. 38:113-136.. 6 pis. naturSagefischen (Pristiophorus M. 1 map (p. 1911. 80. 1905: 97-102. L. 1907. Laboratory and field ecology. Proe. B. 1888. Scrivenor. Stanford Univ. A. Z. Soc. Animal communities of bottom and their importance for marine zoogeography. Studier. 1906. wiss. 1939. Heinrich.. 179-234. 1907." Ibid. Between xxii.. JaJirb. "Environmental studies on the limpet. 13:558-570. App. Williams responses of animals & Wilkins: xii. 73.. Saville-Kent. Shelford.. Baltimore. Semper. "Geological and climatic factors affecting the dMribuProc." Rept. biotic communities of the Monographs. figs. Ges. 15 figs.. G. S. and Jack Calvin. interpretations. mode de formation. 379.. and F. 6 Hill. 54. 12 figs. "Beobachtungen und Betrachtungen bei iiber die Beziehungen der Augen Physiol. 75. V.. 1905. the as indicators of correct working methods. V. 65 text figs. 64. 232 ff. Allen: xviii. Petersen. E. 154:120-165. Press: 69. al. Shelford. 68. "Cours d'oceanographie: les iles coralliennes flore. 71. 11:207- 324.." Verhandl. and Rept. The Bcnthal P. (p. 10 pis. 20:1-76. J.. Reef Comm. zum Nahrungserwerb Fischen.. Ecol. 5:249-354." pi. Richards.. 1893. 70. Scheuring. pt. Freunde Berlin. Oceanog. Stephenson. and C. 1921. Ludwig." 79. Sitzber. text figs. Russell. the sea its food and quantity. Romer. William. Fritz.

3 text figs. Wilhelmi. 84. Verrill. W.. and A. Rept. Malacol." Mitt. 1930. 1930. II. The oceans. 1914: 189-276. "Studies. 11 text figs. their and general biology. Thomson. 1931.S. 1:59-81. 2 89. physics. M. Max. lenger during the year 1873 and early part of the year 1876. Vaughan. on the relation between corals and zooxanthellae. Prentice-Hall: x. Johnson..M. 3 pis. Bur. 88. U. Priifungsanst. expedition 1878-79.. Proc. G. "Studies on the physiology of corals: I. 1878. Weber. W.. "Report upon the invertebrate animals of Vineyard Sound and the adjacent waters. Yonge.. 6 figs.. U. Fisheries. 82. Y. 1:13-57. Entwurf einer biologischen Analyse des Meerwassers... figs.." Rept. Svcrdrup. "Darwinism and malacology.. II. 22. "Die niederlandische Siboga Expedition zur Untersuchung der marinen Fauna und Flora des Indischen Archipels und einige ihrer Rcsultatc. The effect of starvation in light Ibid. "Studies. C. 46:182-191. 8:272-286.. Soc. "Corals and the formation of coral reefs. Sci. The voyage of the Challenger. Harper: 2 Ann. W.. chemistry.. 1942. 38 pis. 90. 1872. No. B. Sci.. 5. N. vols.: II. Y.. 1919." Mitt. Smithsonian Inst." 2 pis." 86. Feeding Great Barrier Reef Exped.: V. 34 text figs. 1900. Yonge. 7 charts. 85. 265 83. 1087. C. etc. and R.Bibliography Nordenskiolels ishafs 271 Svemka Vetenskapsakad. A. C. figs. B. 168 text figs. 1909. Fleming. S. 1:295-778. Wasserversorgung Berlin. mechanisms and food. M. Julius. and darkness 1:177-211. 37 pis." Great Barrier Reef Exped. "Die Macroscopische Faune des Golfes von Neapel. T. vom Staridpunkte der biologischen Analyse des Wassers betrachtet.. Nichols. 1912.. E." Kept. ChalN. M. with an account of the physical characters of the region.. Digestive enzymes with notes on the speed of digestion. Idem. etc.. The Atlantic: a preliminary account of the results of the exploring voyage of //. London. Woodward. Rept. 91. Petermanns 87." Bihang." 6 figs... 16:47-166. Handl. . 43 pis.

This capacity is variously developed. however. In some cases the pelagic period is extremely short. such as the mackerels Scomber scornbrus. Even unquestionably pelagic forms.02 and 1. The lower limit. all daylight had vanished. at 610 m.02 to perhaps averaging about gravity of sea water varies between 1. and varies in each locality according to the time of year. The depth to which light penetrates decreases from the tropics to the polar regions.06.03. many elements in the pelagic fauna. whereas the specific 1. are thought to spend part of the year resting on the bottom. lies between 200 to 600 m. never sink to the bottom. 1. They are not sharply distinguished. for practical purposes. or abyssal zone. The general discussion of the pelagial may best be restricted principally to the lighted zone. although an ill-defined interme- be recognized. like the ctenophores. leaving the abyssal for treatment with the * The common character of all pelagic of the bottom. Off Bermuda on "an exceedingly brilliant" day. Some animals. earlier he had found no visibly detected light below 511 rn. are larvae of animals that spend their adult life in the benthal.04. animals is their sinking. which means that thoy independence have the ability to maintain themselves in the open water without abyssal benthal (Chapter 14).The of the Biotic Divisions Ocean the (Continued}: Pelagial AS IN THE BENTHAL. the lighted and the lightless. Beebe 7 found light still visible to the eye at 571 m. its specific gravity ranges 9 from 1. TWO VERTICAL ZONES MAY BE DISTINGUISHED IN THE pelagic communities.. depending on the angle of diate stage can incidence. lla>3fl Special adapta- 272 . Flotation mechanisms Living matter is heavier than sea water.

as shown by analysis: . This distinguishes pelagic creatures from the animals of the benthal and gives them features in common that appear in various groups by convergent evolution. 54 The shells of pelagic crustaceans are uncalcified or comparison with those of their benthic relatives weakly and have a higher fat content as well.28/* to 18/x.34 Pelagic Anomalwra Fresh-water copepods (Mil 9/21 5. has a very thin shell with walls from 1.5. and likewise with any increase in the resistance offered by the water. and uncalcified.7 calcified in < t Ash <. horn-like. This resistance is the greater. lacks the lime bodies in- variably present in Among and pteropods have delicate shells or none pelagic snails the heteropods The shells of the at all. especially . while the pelagic Loliginidae and Oigopsidae have the shell narrow.13.73 fl. also reduced in the shells of Foraminifera by of the pores and by enlargement of the opening of is the shell.Ol The pelagic sea cucumber Pelagothuria its relatives.5 *. Among pelagic fishes. Globigerina Calcium carbonate increase in the size pachyderma.83 3. The state of suspension may be regarded as a retarded sinking in which the rate approaches zero as a limit. The foraminiferan Orbulina universa from surface waters. The rate of sinking depends on various factors. Reduction in specific gravity may be made in many ways.91 19. whose name indicates its character. delicate.<*7 41.79 *.71 13. of the water.Flotation tions Mechanisms 273 are consequently required to prevent animals from sinking. the more water particles are displaced and the greater the sum of their paths described in such displacement. the benthic Sepiidae have a calcified internal shell. The pelagic Globigerina are distinguished by thin-walled shells from the single benthic species.50 Crangon J/0*i# 3. 38 There may be economy in the use of hard skeletal or shell materials such as lime and silica.. or viscosity. it is reduced with reduction in specific gravity. the cephalopods. Pelagic relatives of benthic animals with a skeleton or shell are accordingly characterized by great reduction in such hard parts. it varies also with the internal friction. 37 Similarly among pelagic lamellibranch Planktomya are uncalcified. while specimens from the bottom have walls up to 24/z in thickness. which depends mainly on temperature and salinity.i Fat Bent hie Littoral Neritic- Nrphmpg Ctircinu* .

are also surroundings without injury since their body fluids are isotonic with sea water. fat. watery and transparent. but the relative is reduced. the pieces passing through the meshes like jelly through a decapod Grimalditeuthis richardi is so transparent that print may be chyma filled The large vacuoles in the parenof pelagic turbellarians of the genus Haplodiscns seem to be read through jelly.5% in a variety of forms. as in Regalecus the skeleton and Crystallogobius. Alciopidae and Tomoptem among Such annelids. Among medusae the amount of water may reach 96. their eggs singly instead of carrying The most widespread means pelagic marine animals is the taking of reducing specific gravity among up of large amounts of water. This water is ordinarily incorporated in the transparent jelly-like tissue. the transparent alberti pus so soft a body that it is cut by the threads of a mollis has Alloposus sieve. 18 Siphono- phores. of fat is widely distributed among pelagic anilowers the general specific gravity. This is not uncommon even among pelagic Protozoa. and the the cellulose mantle is swollen Salpae Pyrosomae. 8 33 The ctenophore Beroe is said to have a fluid of low specific ' gravity in the numerous vacuoles by muscular contraction at need. Reduction of weight is achieved by pelagic copepods by depositing the egg sacks with them. The accumulation and this More extensive accumulation of fat is found in pelagic crustaceans such as cladocerans and copepods (see table on p. 273). which may be emptied mals. Leptocephalus. such as water of less salinity. and the same is true of Noctiluca.274 The Pelagial is among weak swimmers. have a lower specific gravity than sea water. and the cods (among bony fishes) fat bodies are present in the . so abundant among marine animals. Among with water. widespread among the pelagic cephalopods. heteropod and pteropod snails. Numerous mantle of Planktomya ( which may be the larva of a mytilid). or even air. Invertebrate marine animals may take up water from their and the eel larvae. 20 with The plankton fishes. or reduced. The selachians. The and the protoplasm of Noctiluca. ( of its cells ) . More effective than the addition of sea water is the storage of lighter materials. little calcified. It is known as the mesogloea in coelenterates. fluid in the vacuoles of the extracapsular body of radiolarians. Crystallogobius and Aphid. The difference absolute surplus of weight remains the same. its body. the coarse net. Radiolaria contain oil drops in their intracapsular protoplasm. Tremoctojelly resembles a small transparent ball of jelly. weak. is and chaetognath worms have similar jelly-like tissue.

When an excess of density still is present. air sacs ( pneumatophores ) are present. was made possible by the acquisition of an air bladder. so that they may be deep-bodied and even wedge-shaped ventrally. thus their independence from the bottom is The bony fishes are by far the most numerous of the complete. The air bladder has also made the bony fishes more independent in the matter of body form. 17 as the selachians never are. fishes. where no place of rest is available. mackerels. The most effective means is of for reducing the excess of weight the inclusion of air or other gases in the animal body. and seals. serve the same purpose as does the air bladder of that have taken snakes. on the surface. Thick layers of fat enable the basking shark Cetorhinus maximus and the ocean sunfish Mola mola to sun themselves motionless on the surface of the sea. penguins. such as air chamber and jelly tissue in the siphonophores. and fat accumulation and air bladder in the moonfish.Flotation store food material in the fishes are floated Mechanisms fat in their livers. as in numerous Clupeidae. sirenians. seals. The pelagic snail Glaucus (Fig. of fat varies with the season. Similar ac- among warm-blooded marine animals. serve also as insulation. freeing them from the necessity of having flattened lower surfaces and large pectoral fins. such as and The amount whales. One may say that the change from a littoral to a pelagic existence. which usually have an air bladder whose gas content is under control. in addition to their breathing function. An air sac is best developed among bony fishes. vertebrate animals of the pelagial. branched air sacs are connected with the atmosphere and filled with air by rhythmic breathing motions. so that their weight may be exactly adjusted to that of the water displaced. The cephalopods Nautilus and Spirula have a chambered shell with air in the chambers. means of reducing the density may occur in varied combinations. for only the extremely powerful sharks and a few types of bony fishes without an air bladder are capable of the unceasing muscular exertion necessary to keep from sinking. it must be overcome by increased These different . which are In Velella. and in non-polar regions it is usually true that there is cumulations of fat more fat in the warmer season when the supporting all power of the water is least. and flatfishes. Among siphonophores. the lungs. 275 form of The eggs of many by the inclusion of large oil drops. which floats filled with gas from a gas-producing gland. Among the air-breathing vertebrates up a marine existence. 530) has intestinal gases that are sup- posed to play a similar part. such as sea turtles and certain and whales.

Spence Bate. Phyllosoma larva of Palinurus. X 18. d. f. g. d. 53. Tomopteris euchaeta. Maldanid larva ("Mitraria miilleri"). Glaucus c. g. Larva of Sergestes ( ElaphoAfter Regne Animale. X X e. Suspended or atlanticus. b. slightly enlarged. a. 40. natural size. b. e. Chun. . Agassiz. and Giesbrecht. Calocalanus pavo. floating pelagic organisms with enlarged surfaces. a. c. Larva of Lophius ) piscatorius. caris dohrni X 13. Haecker. 2%.276 The Pelagial FIG. /. somewhat enlarged.

The amount of resistance in the water. Such motion may consist . 56) have relatively long tentacles and ambulacral free-floating life possible until their increase in outruns their weight enlargement of surface. This means of support is less developed among the Metazoa. crustaceans. Such apbler. Lcptodiscus. which increases with the increase of the horizontal surface. Young transformed starfishes and sea urchins (Fig. a pelagic forflattened. paratus is most developed among the Radiolaria and Foraminifera. intrinsically The high surface-weight proportion.Flotation friction Mechanisms 277 with the water and resistance offered by the surface. which make a appears only among small and very small animals. often occur. depends on the sum of the distances to which the water particles are forced when the body is moved. Such "suspensory bristles" are notable in Hastigerina (Fig. Lateral projections. The is by changes in form a widespread phenomenon among pelagic inretardation of sinking It is possible to equalize only a small excess of weight in this manner and it is effective only for small animals. the FIG. 55). 53. including annelid worms. are shown in Fig. which have an vertebrates. 54) and Globigerina (Fig. The great majority of pelagic animals exist without this means of suspension. One may speak in this sense of the form resistance of a body. 53d). simplest means of enlarging the ventral surface lies in the flattening of the body. in which the long pseudopodia are supplemented by spines of the test. and a fish larva. the crustacean larva Phyllosoma (Fig. aside from friction. Such resist- ance to sinking may be actively increased by swimming. the pelagic nemertines. which may be aminifer. the light-producing flagellate. prodpced by active swimming. some of the most conspicuous examples. and many others are thus digitata. it feet. Water resistance. mollusks. is the most widespread means of preventing sinking. the turbel- larian Haplodiscus. 54. their larvae. In general this increase of surface by the development of projections is not widespread. Haxtigerina copepod Sapphirina. most Hydromedusae and Scyphomedusae. The amount of such displacement is increased with increase in size in the horizontal plane of a sinking body. After Rhumbranched in addition. A sheet of any material will sink more slowly if laid in the water on its side than if placed edgewise.

278 The Pdagial largely of force exerted opposite the direction of gravitational pull. p. pedicellaria. 56. FIG. or may be a small component of lateral motion. spines. After Zicgfar. 55. as in the pteropods. Juvenile sea urchin Arbacia pustulosa: f. as in the sharks. its The water body is proportional to projection on a plane at right angles . After Korschelt and Heider. a pelagic foraminifcr. FIG. ambulacral feet. Globigerina btittoides. st. The effectiveness of the reaction its is measured by resistance the size of the of a body and rapidity of motion.

After Mortensen. In the almost microscopically small larvae of echinoderms. and among these especially in larval stages. Cilia and lashing hairs. the largest has the relatively longest ciliary band.Flotation to the line of motion. worms. they are arranged in narrow rings or bands. very effective .. 57.and are found mainly among protozoans. on account of their small surface. Mechanisms 279 speed with which the and to the square of the body moves. Auricularia larvae with different lengths of larval life. Balanoglossus. further support can be gained only by rela- . among very small Metazoa. When the larvae increase in size. etc. Great numbers of cilia are required to support even small animals. X about 35 times. are not FIG.

The larvae of echinoderms rarely reach any considerable size swiinming stage. has a of turbellarian diameter Haplodiscus disk-shaped pelagic only 1 mm. Pelagic cephalopods and rays swim by the undulation of lateral fins. The contractile bodies of salpas and pyrosomas are more or less tube-shaped. Among pelagic annelids the parapodia are employed for this purpose This form of locomotion is outswim and catch fishes 1 number (Alciopidae. the second pair of antennae fulfills this function and may be much enlarged. Sea turtles and penguins row with their forelimbs.280 The Pelagial tively great increase in the length of the bands of cilia. but in those that do. which on which they feed. water resistance is uniformly pro- swimming duced by muscular work. but even so this device is suitable only for small animals. In pelagic crustaceans.. Fig. It has been convergently developed among pelagic cephalopods by web-like membranes connecting the arms (Arnphitrctus. and another turbellarian in the North Sea. to which. in the former. 58). in length. In the heteropod snails . respectively. Undulation of the body is the most common mode of swimming. in small forms like the copepods ( Fig. forms chains 0. the ciliary bands are folded and bent in a remarkable way (Fig. 53c). The bell shape ejected prevails among the Hydromedusae and Scyphomeclusae and in the swimming bells of siphonophores. Tomopteris. Ahnirina.5 mm. Cirrothauma. of water. and among pelagic holothurians by similar webs between the tentacles (Fig.9 to 2. Fig. The means used method consists in the production of a stream striction of a vary. A more favorable arrangement is the distribution of the cilia over the whole body surThe face. 1 " Among the pteropods two parts of the foot are transformed into oars. in Sergestes and Polybius. The pelagic decapod crustaceans are distinguished from their benthic relatives by the broadening of two or four pairs of posterior thoracic limbs as. highly developed by the squids. 60 ) and small larvae. is added the darting produced by sudden expulsion of water by the muscular mantle. whose eight rows of during their freeplates are composed of fused cilia of relatively great length. 59). Ctenophores. Oar-shaped jointed limbs are a frequent form of swimming organ. the bipinnarians and auricularians. 57).. Among larger pelagic animals. are forms with extreme development of gelatinous tissue. numerous limbs may be employed for swimming. among schizopods and numerous pelagic amphipods. a varying of limbs are used. reaching a length of about 5 mm. A widespread by the con- tube or bell-shaped hollow body. the reaction from the water driving the animal forward and upward.

swiniining membrane between the ampullae. After Ludwig. a pelagic holothurian. wi. 59. mouth. ampullae. . f. a pelagic octopod.. After Chun FK. Cirrothnuma murnnji. anus. Pelagothuria natans. . d. 58.Flotation Mechanisms 281 FIG. tentacle. r.

whales. porpoises. is suspension. In most fishes. X Brady. Swimming. The cnpepod Cdlanus at the left. may be defined variously combined with other means of suspenas distinguished from motions that merely produce as motion sufficiently active to render the animal independent of the oceanic currents. as a rule. Such swimming is never produced by cilia or by vibratory motion. Active motion sion.. The effectiveness of muscular motion is such that other arrangeis and Small animals less ments for suspension are rarely combined with it. has several groups. with their depressed bodies. ward. perhaps form an exception. bony fishes without air bladders. 60. This increases proportionately with the size of the animal animals. long (and hence all microscopic animals ) are incapable of such swimming. in larger than 1 cm. among . in these vergence been attained by con- Fie. but is always dependent on more typical muscular exertion. accordingly greater. The resistance to the waves produced by these motions drives the fish forforce of gravity. St. with its bristles. The antenna used as an oar finmarchicus. and seals. After (greatly enlarged). at the right. the maxilla 26. small free-swimming animals the suspensory processes are placed in . For the rest. and the component acting on the lower surface neutralizes the Thus a special type of body form in large pelagic sharks. and seals the whole body or the posterior part of the body undulates. almost round with a slight ventral flattening.282 The Pelagial a median ventral fin is undulated. The large pelagic rays (Mobula and Mania) reaching a breadth of 7 m.

and all microscopic animals that are pelagic belong to the plankton. but they merit discussion as including assemblages of animals with much in common. Most of them are small with parison or microscopic. Plankton includes all those animals and plants that drift aimlessly. The giant i. which reaches a diameter of 2 m. . pension with that of a rudder. water even small animals may be capable of directed movements. bitrary. is whose independent movements are insignificant in comthe movement of the water. such as the Cyanea arctica.Plankton and Nekton 283 the plane of motion (for example. Some modern students use the term seston to apply to all swimming or floating bodies in the water. Complete independence of currents dependent of wind and some cephalopods. 53g). the free-swimming forms are called nekton as disis tinguished from the passively floating or suspended plankton. thus they do not obstruct the motion and combine the function of aiding sus43 44 In the swiftest of the swimmers. the nonliving seston then called tripton. Both plankNekton is tiles.) The converse is not true. * Plankton and nekton Pelagic animals freely. The definition of these several sizes of centrifuging. the living seston associated intimately with the surface film is called neuston. or according to their dependence on the latter. alive or dead. The plankton. until it was discovered plankton -escaped observation It is now secured apparatus of the appendiculates. may be grouped according to their ability to swim independent of oceanic currents. such as a nocturnal rise to the surface.7~fHose jellyfish shark Cetorhinus niaximus is so very dependent on the Atlantic curIn quiet rents that it might almost be included with the plankton. such as the blue shark and the mackerels. own their net V PlaiSton the by escape animals ' and megaloplankton. A especially vertical ones. minute nannoin the feeding by filtration or is ar- plankton which are mainly incomposed of free-swimming forms. It evident that no sharp division between these groups exists. mesoplankton. in Calocalanus.e. are unable to of plankton includes those forms that practical definition movements. ordinary plankton micromay be assorted according to size as nannoplankton. the marine repis reached only by certain fishes. Fig. as many larger animals with muscular movement are also included in this category. and in the mackerels the body fins fold into grooves when not in use. and the homoiothermal marine birds and mammals. current. every unevenness of the body is removed.

salpas. for example. Challengeria xiphodon " swirei 0. the smaller ones in the warmer sur- warm face waters. which is of such great importance in reducing the rate of sinking.28 warm The same mantha. The arrowworm Sagitta Metazoa hexaptera.11 1 may be grouped 400-1500 in. from 1500-m. It depends chiefly on the "temperature. Other conditions being equal. only specialized forms like Mold with its thick layer of fat. " " slogetti " " 0. depth averaging 30 mm. " " PharyngeUa gaxtrula 0. Such feather-like processes as are found in Calocalanus (Fig.284 The Pelagial ton and nekton include predaceous as well as herbivorous forms and cannot be distinguished according to their food habits. specimens from 500-m. with its and sub- siphonophores. less markedly on the salinity of the water. which inhabits the lighted pelagial. 60 mm. medusae. a sinking of the animals which they can maintain themselves. 3 fish Cyclothone microdon increases in size with depth.38 Ch. like the sharks.%15 " mm. is not uniform. The viscosity of water. " Ch. it is only half at 25 what it is at 0. Among the pelagic cephalopods the smaller forms are found only at the surface. 0. other fishes are represented mostly by juveniles. Pelagic dinoflagellates like Ceratium form longer processes in warm than in cold water.58 " hantoni 0. bethelliQ..1 mm. chanical relations with depth throw light upon the occurrence of sur- . are found at the Such mesurface. Ability to float seems to be the governing factor in the composition of the surface fauna of the tropical tropical Atlantic. and pyrosomas.. 53g) occur in warm-water copepods but not in polar forms. Among larger fishes. while the larger indent the oceanic depths. is larger The little and more mature the deeper the source of specimens. according to depth: 1500-5000 in. 61) are assorted vertically according to size. the larger ones below. or active swimmers. for example. The radiolarians of the family Challengeridae (Fig.85 0. There is similar sifting by to the depths at -' - size in the shrimp Acanthephyra..16 tizardi 0. Aulacantha scolysifting.5 0. The conditions for suspension of organisms are consequently more favorable in the cold waters of the polar regions and of the oceanic depths than in the tropical waters. is This seems like a mechanical exhibit the true of other radiolarians. Numerous same phenomenon. and specimens from the same depth in the southern part of the North Atlantic are much smaller than the northern ones. thomsoni u mm. Thus the average sizes in milli1S meters of the following forms 50-400 m.

they occur: Various sizes of Challengeria species according to the depths at which 140A. slogetti. C. such as the pteropod Clione.Plankton and Nekton face forms in the 285 Norwegian seas. where the viscosity is the same as that of the surface waters near Norway. C/i. in the warmer part of the Atlantic. After Haecker. . naresi. Challengeria xiphodon. 30 The variation of viscosity with temperature may bear upon the fact that many pelagic animals are smaller in warm seas than in cold. and the copepod Calanus. X depths of 750 to 1000 m. the medusa Aglantha. C/i. H. 61. which occur also at FIG.

convergently evolved from different origins into similar structures with pelagic animals.. The larger plankton animals. and there are none of the many-celled algae and vascular plants. signata in the vicinity of the Canary Islands are notably smaller than at the same depths in the North Atlantic.) serve the same purpose. This matter evidently requires consideration in connection with the size relations discussed ^ in Chapter 10. and the arrowworm Sagitta bipunctata reaches a length of 12 mm. which i minute compared with themselves. Lohmann found. as it applies also to the benthic animals.5 The devourers of these plankton algae are in part small ani- mals. Foraminifera. but they also include larger ones that have solved the problem of securing large numbers of the small forms. The long tentacles of siphonophores (up to 30 m. like functions. 60). The fishes Cyclo- may be thone microdon and C. those that produce a current of water.. and those that hunt actively. that for every metazoan (such as an ephyra or Sagitta) there were 1000 protozoan and 7000 protophytan cells. in the Mediterranean and of 44 mm.) The basic food supply is the plant portion of the* plankton.286 This The Pelagiat referable to direct action of the lower temperature. such as radiolarians. small Metazoa up to the size of pteropods. the . divides plankton feeders into three groups: those that feed by means of tentacles. from 21 which the food is removed by the is specially adapted in animals that lips. but it involves nonetheless a coincident adaptation on the part of the pelagic fauna. form the food of the larger In order to secure their food. the plankton feeders require special apparatus. in the Bay of Kiel. such as are found nannoplankton of the ocean. that endless number of single-celled algae. The into Chiroteuthis has its disks evolved cephalopod sucking sticky threads. since their food differs in a number of ways from that of benthic There are no great accumulations of detritus such as occur types. A most singular means of securing food found strain out the living forms from a stream of water that they produce themselves. which paralyze their minute food by a poison. The bristles of the mouth parts of Cladocera and copepods (Fig. etc. The tentacle feeders are the most primitive rest. and these figures are in the low. they feel about for their food while otherwise remaining at Radiolaria and Foraminifera do this with their radiating pseudo- pods. in the Arctic. Special means of securing food are required by pelagic animals. Lohmann 24 type. Self -produced nets or screens perform this function. copepods. on the sea bottom.

Predaceous animals. The groups bryozoans. most ctenophores. this all illustrate this is apparatus means shown by the of food-getting. left The appendiculate Oikopleura is albicans (black) in its house. and With the exception of the Chaetognatha. heteropods. siphonophores. 62. The black arrows show the direction of the currents of water produced by the undulation of the tail. pteropods. ascidians. and snails only .Pelagic slime bands in the and Benthic Formations Compared body cavities of salpas 287 and pyrosomas. comall exclusively pelagic. do not present any struc- tures peculiar to the pelagial. salpas. Lamellibranchs are represented only by a few opisthobranchiates and the and pteropods. the screen-like gill strainers of the plankton-eating fishes. which hunt for their prey by active movement and find it by means of their sensory organs. To the the permanently outspread net. and appendiculates are The copepod crustaceans predominate. worms are few. and the baleen of the toothless whales. The of the animal light arrow beneath points in the direction in which the animal moves. chaetognath worms. absent in the pelagial are the sponges. Radiolarians. echinoderms are repre- sented by Pelagothuria and larval stages. brachiopods. After Lohmann. was discovered by examination of the FIG. the nannoplankton. forms.895 small copepods. built up out of jelly-threads (Fig. Pelagic and benthic formations compared The special adaptations required by the pelagial give its fauna a somewhat different composition from that of the benthal. 62). and Large groups are represented by only a few specialized There are a few pelagic actinians. the remark- able apparatus of the appendiculates. which had escaped the finest silk nets. medusae (with specialized heteropods the exception of Lucernaria). by Planktomya. such as the herring or the giant sharks. appendiculate sieve and by the presence in the stomach of a single herring of 60. The effectiveness of fact already stated that a whole new world of minute plankton creatures.

The same true of most species of the pteropod genera Hijalaca and Clcodora. i. and snails. neither fortuitous nor uniform. and chaetognath worms. Distribution of pelagic animals fishes are also an important ele- The free suspension of pelagic animals favors the wide distribution. 25 Long-lived animals can exist for periods of time in unfavorable localities. cephalopods and ment. No pelagic forms appear to have been discovered in one ocean that are not represented by parallel forms in the other. but also of many with poor powers of swimming. The Acanthometridae (Radiolaria) are astonishingly similar in the warmer 3The siphonophores of the parts of the oceans of both hemispheres. but ecologic. Its theless.. siphonophores. But as these are also ultimately dependent on the plant plankton. dependEach smallest portion of the ocean all times a population of nannoplankton and microplankton that an exact reflection of the environmental conditions. as in has at is polar seas. annelids. corals. and hyperine forms in addition. which occur in all the oceans. sea anemones. the animals are abundant . distribution is ent on environmental conditions. varying from 20 to 60 days in the echinoderms to 4 to 7 days in brachiopods. bonito. and of the heteropods Atlanta peroni and Oxijgyrus keraiidreni. which have a pelagic trochophore larva. and others. such as the deep-sea fish Ccratias couesii. Many planktonic forms have the same distribution. and mackerel. drawing upon their stored supplies. decapod. many benthic animals is of great The length of larval life becomes a governing factor in the extent of the distribution of such forms. are more widely spread than Melo and Voluta. The short-lived and rapidly multiplying nannoplankton affords the best index to the existing conditions.288 The Pclagial posing 90S! of the whole fauna. Conns. Never- and pelagic life is accordingly not uniformly distributed. with schizopod. Most of the important genera of sharks are found in both the Atlantic and Indo-Pacific. and Cypraea. The pelagic life of the larvae of importance to their distribution. from whose large-yolked eggs the young snails appear in a benthic juvenile stage. Hedley has shown that in Polynesia the gastropod genera Mitra. the conditions are not uniform. not only of active swimmers like tuna.e. two oceans are often distinguished only by trivial characters. Variations in environmental conditions are less frequent and less abrupt in the open sea than in the shore waters or on land. Next come the pteropods. pelagic is turbellarians are represented in both by the same species.

25 Among longer-lived animals. in which animals of varied kinds are so abundant that one can dip a soup of plankton. cod. Salpas and pyrosomas. The composition of the plankton varies with time as well as with It is usually composed of a great number of different anilocality. mals. The extremes of shown in the poorest catch of plank- 763 organisms per a maximum -6 liter in the tropical Atlantic. divided the pelagial into rich and poor sections according to the existence of more or less than 1000 nannoplanktonic organisms per liter. 1 appear in the harbor of Messina." the latter as "swarm" aggregations. ctenophores and siphonophores.Distribution of Pelagic Animals 289 where the plankton ton. and the impoverishment of the plankton begins below this level. as in the schools of herring. of the sea water. since the plant element in it is dependent on light. The basis of study of plankton distribution in this direction lies in the construction of lines of equal plankton development or "isoplankts. there are vast aggregations. for example. of pelagic distribution on this basis has just begun. and developed the more accidental accumulations on the borders of currents and in The former have been distinguished by Apstein as "producbays. So-called "animal streams" 14 rent. 60) may color wide stretches of the sea brown. Such an accumulation observed by the German Plankton expedition in the Atlantic extended for 300 km. and other fishes wandering to feedSuch aggregations also appear in ing grounds or breeding places. such as the annually in July may appear in both open sea and near sometimes with a good deal of regularity. northern copepod Calanus finmarchicus (Fig. but at a given place and special time a single species may flour' 42 plankton meet. mackerel. with a breadth of 5 to 10 m. In the rich areas the distribution is not uniform. twice daily." especially for equal amounts of the plankton as a whole and for certain out- standing species. These streams may be distinguished by a smooth oily appearance of the surface and may extend for more than a kilometer. The great numbers of the tion. numbers of Salpa fusiformis that are and August north of the Hebrides. and even globigerinas are found in great masses. is density of life in the pelagial are at least periodically rich. Their appearance may be dependent on wind and curcoasts.915 in a liter in the cold water of the North Atlantic. The siphonophore Vclella forms swarms of enormous extent. they . The study Lohmann numerous distinct aggregations of The rich domain of pelagic plankton as especial density of life appear. as compared with of 76. is the upper 100 m. especially in places where currents rich in less active forms.

Such plankton may be dominated either by mature or predominates. The maxima and minima of different organisms depend in different ways on the temperature and may alternate in the single elements so that a varied plankton Fie. to return before daybreak.290 ish to The such an extent that it Pelagial nous plankton. one may distinbetween The oceanic these as oceanic and neritic pelagial. This is the North Sea a pteropod or copepod. to permit comparative studies Perhaps/ in time. After Lohmunn. To this group belong all radiolarians. even from great depths. 63. 63). in the Bay of Kiel. . at different seasons (Fig. these animals are called holopelagic or holoplanktonic. Schools of herring come to the surface at night. guish life is made up of animals of the bottom pelagic independent throughpelagial of the out their development. At night numerous deep-sea forms come to the surface. at Curves of volume of various groups of organisms in the total plankton Laboe. during the year. These are followed by predatory forms that live on them. and in warm seas a salpa. Important and readily understandable differences exist between the deep sea and that of shallow seas. may predominate. There is even a difference in the composition of the surface plank- ton between day and night. in by ton larval animals. but remain at considerable depths during the day. such plankton com- may have very different compositions Changes may take place too rapidly by one ocean-going vessel. parisons in distant regions may be made by cooperative effort or by the use of adequately equipped airplane expeditions. many Foraminifera. and one speaks of monotoespecially a phenomenon of shallow seas. is important in Seasonal variation in composition of the planktemperate and cold seas.

velellas. and currents carry the neritic forms out and the oceanic ones in. appendiculates. copepods and hyperines among salpas. which comes to the surface in and some without. such as salpas and In the spring of 1900. a great number of bony fishes with The mals the terms hemipelagic or meroplanktonic are applied. Windrows of dead velellas more than half a meter high and a kiloneritic meter in length appear on the coast of the Riviera after storms. which feeds on benthic animals. and finally the Cetacea. The water fleas of the genera Podon and Evadne. air bladders. etc. 22 Both oceanic and medusae. fiords. among the latter Pelagia noctiluca).Distribution of Pelagic Animals the 291 medusae without a sessile stage (many Hydromedusae but few Scyphomedusae. anidependent on the bottom during part of their existence. plankton suffer in storms and are driven ashore in great masses. to these cludes rays. Finally. To this group belong all the medusae with alternating generations. whose eggs sink to the bottom. such as greater turbidity of the water. in Deep bays. the larvae of numberless benthonic animals. in addition to holopelagic forms. and especially the walrus. contour. so that periodic changes in the composition of the neritic pelagial are much more pronounced than in the oceanic. even some active swimmers like Julis and Com. which rest on the bottom at night. operate like which large numbers of macroplankton. Hemipelagic life infor example). The inhabitants of the neritic and oceanic pelagial are mixed at the boundary. The time as well as the season of dependence on the bottom vary with the different animals. May and many bony fishes. neritic pelagial includes.. the neritic pelagial includes the sea turtles and seals. This boundary waters. lagoons of coral reefs. generally To to their destruction. and the floating eggs of many fishes. most sharks and cephalopods (Sepiola. and the ostracod Philomedes brenda. and ianthinas had been driven into the Bay of Naples in such numbers that they covered millions of square meters. physalias. are hemipelagic. pyrosomas. many to breed. the chaetognath worms. the obvious result of dependence on the bottom. in the neritic area and to what extent they are destroyed by biotic competition is not known. accumulate at certain seasons. most siphonophores and ctenophores. some sharks. although the eels are a marked exception. in . heteropods and pteropods. traps. all were stranded and destroyed by the beginning of May. crustaceans. what extent the holopelagic animals are destroyed by modified physical conditions. is The is neritic pelagial bounded in general and thus includes banks as well as coastal by the 200-m.

Sweden. The food chain here represented extends from the small single-celled algae to the larger fish and toothed whales. which is able to reach great depths (up to 750 m. adapt themselves to the change. corresponding to variations in the plankton.). The dependence of various elements of the food chain on a preceding one conditions the distribution of the larger forms. plankton. The baleen whales are at home in and antarctic seas. which have been able the neritic area. 10 aside from their breeding movements. and the richer fertilization of the water in the Atlantic. in The number meter of water may of appendiculates in a tenth of a cubic reach 600 in shallow water. warm seas the plankton lies at a deeper level and is followed by large crustaceans and cephalopods. dominated by food supply.292 August 1883. are found in the stomachs of the sperm whale Physeter macrocephalus. The population of the pelagial is a well-characterized association almost entirely self-sustaining. the differences between them being greater than that between the polar and tropical food supply provided by coastal plant detritus. it was good in 1905. directly. harbor the humpbacked whales Megaptera. and good again is in 1907. The coastal waters. and neritic ones rich. in turn. The mackerel fishery at the mouth of the English Chan- depends on the amount of animal plankton. rich in plankton and also the arctic small fishes. are probably a response to the The distribution of whalos f is distribution of their food supply. while it sinks to 13 in the waiter of the 23 open ocean. Gulmar Fjord. The complicated wanderings of the herring. the beach The Pelagial at Kristineberg. the holopelagic copepod Anomalocera patersoni. Plankton bacteria are which make possible a greater development of plant much less abundant in the open ocean than near land. was and in covered with a thick glowing mass of the protozoan Noctiluca of broth at that thick water was a the place greenish September 1893. Oceanic areas are poor in life. especially copepods. lies 25 The reason in the better vicinity of land. the north coast of Iceland of the The appearance of the herring on dependent on the summer development copepod Calanus. and hence rather sharply distinguished from neighboring associations. . In open. Giant squids. It is increase in the animal population follows the richer food supply that keeps various holopelagic to An animals. as already shown. where such an excess of plankton develops at certain seasons. poor nel in 1906.

farther north. they apparently winter in deep water and re-enter the circuit at the Faeroes in spring." colithophorids. The latter carries water from the neighborhood of the equator to the 48th parallel of south latitude. animals to the same position and enThe distribution of the developmental stages of Calanus its 12 Great numfinmarchicus in the Norwegian Sea affords an example. and the currents of the depths are only beginning to be studied with reference to their special faunae and effects on distribution. the larvae emerging from these eggs gradually grow.Pelagic Biotopes Pelagic biotopes 293 Distinct biotopes within the great extent of the pelagial are only vaguely differentiated. This is still more strikingly a requirement in the great closed circuits of the North and South Atlantic. several months and even years to return to its origin. while the adults die out. current to The may come A closed current returns vironment. bers of adult animals with eggs appear in spring in the diatom beds near the Shetland and Faeroes islands. At the present time only the surface currents are well known. These animals thus are carried through an extensive temperature range and must be to a considerable degree eurythermal. Lohmann this cycle. The action of the currents differs according to their course. are included in this period. especially in its peripheral portion. carried away by flow of the Gulf Stream. and zooflagellates with the seasonal forms in fresh-water animals. and the great oceanic water masses. circuit. Among short-lived plankton animals many generations. are exposed to a considerable range The plankton in such a current requires of temperature variation. or may return to its source. about 1% years in the North Atlantic Current and 2% in the South Atlantic. Larger animals with longer periods of development are also exposed to considerable differences in 25 believes that regular differences. even hundreds of generations. and the number of juniors lessens as that of adults increases. peridinians. . successive generations. when the warm water of the Gulf Stream brings them to sexual maturity. Perhaps the most important differences are those between the oceanic currents. into "juniors". and organisms in it. Oceanic currents are obviously of primary importance in the dis- tribution of pelagic animals. These relations require further investigation. corresponding in cocwith are connected "cyclomorphoses. forming a closed an end. the current then returns from Jan Mayen. and many free-swimming forms are also influenced by them. these then develop. the eddying areas.

The life of the Labrador Current differs from that of the adjacent Gulf Stream. as on the are carried England Banks.294 The Pelagial conditions are quite different for organisms in the non-circulating currents. are unable to breed. On many banks there may be rapid and extensive displacements of water and consequent extreme fluctuations of temperature. so that index forms of cold water such as Oikopleura labradoriensis and Frittilaria borealis reach the North Sea as in spring. or cold water into warm. Such massive destruction provides food for the bottom fauna. is sharply distinguished from the adjacent north and south equatorial currents by vast amounts of the schizophycean Trichodesmium and by the appearance of neritic forms like the appendiculate Oikopleura dioica and the peridinean Prosocentrum micans. the more the current differs in physical and chemical characters from the neighboring waters. so that its composition changes with the Such currents have a characteristic fauna. and. the Agulhas Bank. varying with their origin. caused by change in the relative predominance of the Gulf Stream and Labrador currents. The contrasts are especially notable where a warm and cold current meet. An index form for the Polar Current north of Iceland is the copepod Calanus hyperboreus. which did not reappear for many years. like The The distance the Labrador Current or the Humboldt Current. as does the Gulf Stream or the Japan Current. The plankton carried along by the current becomes mixed with that from other sources. whereas in the Norprogress of the current. The Guinea Current. which either carry warm water into a cold region. 19 and the southern forms of plankton formerly abundant in that area on the border of the Gulf Stream also disappeared. waters over the warmer southern New England Banks in 1882 caused at the juncture of the New extensive destruction of the tilefish Lopholatilus chamaeleonticeps. or Japan Current and the cold Ojashio on the east coast of Japan. Murray showed that the deposits of pelagic Foraminifera on the sea bottom were greatest where currents of difUnusual westward extension of the cold ferent temperature met. and ultimately die. which flows from west to east near the equator.5 between two successive tides in Narrangansett Bay. There may be a great mortality in the plankton at such places. whereas in the summer warm-water forms such 11 to the Lofoten Islands. and are the better defined. Physophora hydrostatica These foreigners suffer in the new environment. Fewkes l3 noted a variation of 5. reached depends on the season. the Spitsbergen Bank. by its disintegration. . for pelagic life wherever the resulting food substances are brought to the lighted zone.

or care Some of these forms. to live a littoral life on the high seas. provision for the young by nest-building. cloud of Salpa flagellifera 100 m. Such pelagial. and associated wealth of life in a some 200 miles southeast of Cocos Island in the eastern off Algae provided with floating bladders are carried by the oceanic currents and accumulate in the quiet eddies in great amounts. especially durhurricane season. is the basking shark Cetorhinus maximus. 2 The Sargassum comes from the coasts of the Caribbean islands is carried away by the strong Florida Current. 2 polychaetes. Various adaptations are necessary. Others appear to have become specifically transformed (such as the shrimp Latreutes ensiferus) and are confined to the Sargasso Sea. 8 4 30 and certain stations on the borders of great currents are especially rich in pteropods. 39 Beebe 5 decarried into quiet bays ' ' A scribes masses of floating debris current rip Pacific. without reproduc- and the bottom. and. . rarely found outside its limits. such as attachment to the weed. A characteristic animal of the North Atlantic Current. like the small crab histrio. long has been encountered on the border of the Benguela Current. the shrimp Leander tenuicornis is also found in the Pacific. The weed is carried back and by wind and may reenter the current for a second accumulations of seaweed form a special biotope in the forth circuit. ing.A wegian Sea Sargassum Community 295 it is C. finmarchicus. The masses of Sargassum are separated out the ing to the right of the current. 2 bryozoans. are common on the Gulf Coast. in the Indo-Pacific there are other species of Sargassum. are widespread on floating bodies of every kind. 31 The change to this half pelagic existence being impossible for some of the original inhabitants of the seaweed. but enabled by it free-swimming forms. just as debris is may accumulate at the borders of cur- by flood water in streams. until the bladder-like floats disintegrate and the weed sinks to The weed affords a base for a curious nomadic fauna 41 coming from the original home of the Sargassum. directly poor in species. and in the South Pacific Macrocystis pyrifera. Planes minutus. Great numbers of animals rents. in the more of littoral origin. its fauna in the Sargasso Sea is Besides a few species of algae there are 16 animals to attached the weed. In the eddy of the North Atlantic Current the weed is Sargassum bacciferum. A Sargassum community The best-known example of such an eddy is the Atlantic Sargasso Sea. 10 hydroids. Others. such as the fish Pterophryne of the young. and continue to grow.

64). A small sessile ascidian (Diplo- The most frequent The nudisoma) . This ( is true of the fishes pipefish especially and sea horse. of unexplained faunal parallels exist between the fauna of this area and that of the Mediterranean. The number These of individuals of the Sargasso animals is fairly large. The Sargasso fish Ptcrophryne makes a nest among the "leaves" by cementing them together with mucus. 64. branch Scyttaea pelagica creeps over the weed. other species with sim- . they may have remarkable weed-like appendages. of the Sargasso animals are similarly white-spotted on a yellowish brown ground color. tubes of the annelid hydroids are Clytia and Laomedea. which this coloration may protect them are doubtless the against 41 sharp-sighted birds. Clytia. and in addition to these such small forms and fishes live in or rest upon the weed. similar radiolarians. Spirorbis.is present. 1 ascidian. covered with the hydroids Aglaophenia FIG. and the weak- swimming crab Planes uses it minuttis as a resting place. besides Pterophryne) Floating piece of sargassum. The weed is yellowish brown. snails. the and the bryozoan Membranitube building worm After Hentschel. with Lithoptera only from these two regions. and the nudibranch. such as similar compositions A number of the fenestrata known nannoplankton. lflrt The Pelagial as turbellarians. the spotted with white by Membranipora and Many Spirorbis. 30 tain crustaceans. as well as cer- The enemies and pora. animals are wholly de- pendent on the weed and constitute a pseudobenthos (Fig. as are the spiral calcareous Spirorbis. The bryozoan Membranipora is abundant.296 1 cirripede.

1947. depth as compared with 33 m. At 200 m. of North Carolina Press: xvii. Idem. Zool Zentralbl. 1934. 16 about 4 higher than in corresponding depths in the adjacent waters. "Biologische und faunistische Untersuchungen an Radiolarien und anderen pelagischen Tieren." Brandt. 24 ff. 1 map (p. LO. 2. 1892. 7. the Secchi disk is visible to 66 m. the Sargasso Sea is characterized by poverty of plankIts water is remarkably clear. 152 and 196 ff. 9. Society. 69 pis. Idem.)." Verhandl.. 80:495-496. "Salpen der deutschen Tiefsee Expedition. Apstein. 1906.. Bullen. 3 pis. This great and wide sea.. R. D. Idem. 16:37. This is explained by the fact that on account of its greater density surface water is here being carried down to the oceanic depths. Putnams.. illus. 23:226-232. and this also explains the poverty of life in general in the Sar- gasso Sea (Chapter 11). G. avec une carte" [abstract]. "Exploration of the deep sea. 8.). 76:344.). 1909. compensating in part for the upwelling currents on leeward coasts. 1926. "Das Tierleben der Hochsee. ton. Idem. deut.. 1906. . Y. E. Bresslau. Tiefsee 4. 1897. in the Mediterranean.. Beebe. in the North Sea. 439. Syst. E. pod Copilia mediterranea. Damas. 5.Bibliography liar 297 and the cope- distribution include the annelid Alciope contrainii. Ges. William. Carl." Exped.. sketch Bull. (pp. "General Agassiz. Nagele: 1-64." Science. Coker. Albatross from February to May 1891. 1901-1903. 325. at 400 m. "Ober das spezifische Gewicht des Protoplasmas und die 6. BIBLIOGRAPHY 1.. E. N. xix. "Oceanographical work at Bermuda of the New York Zoological Science. Alexander. 1-115. Stuttgart. of the expedition of the Museum Comp. the temperature is 18. 1905. Zool. 20 m. Zool. 8ff.). 23:1-89. 1895.. D. Die Salpen der deutschen Siidpolar Expedition. For the rest. (p." C. The Arcturus adventure. and 13 m. 9:155-203. 1913. Chapel Hill: Univ. ton. Chun. "Notes biologiques sur les coppodes de la mer norvgienne. Kiel: Ergeb. Among the few abundant forms are the water flea Evadne spinifera and the larvae of the forms living an the weed. 15 text figs. Wimperkraft der Turbellarien und Infusorien. 14:776-779. Karl.. 174 figs. Llfl. Zool. 9:27-74 (p. 1932. 1907." Reisebegleiter fur See- fahrer. 12:245-290. 3. Zentralbl.. "Plancton studies in relation to the western mackerelfishery" [abstract]." Zool. in the Baltic. 52 ff. 12. The high temperature of the water at considerable depths is a notable characteristic. Jahrb. Beziehungen zwischen arktischen und antarktischen Plank- LI. 7 pis.

20:94-95. Fischer: d. "Observations Troisieme note. 1922. 671). Heft EC. based largely on the scientific researches of the Norwegian steamer Michael Sars in the North Atlantic. 3:1-617. Paris. 1896-1900.. Bioniologie. 1:97-114.. 40 Hansen. Michaelsen. 1-6. 1888. 43 ff. J. Idem. 38. The invertebrates: Protozoa through Ctenophora.. "On certain medusae from Zool. Kiel (n. 6 figs. J. Haeckel. (p." Ibid. 27." text figs. Danish Ingolf Exped. "Tiefsee Radiolarien. La vie dans les oceans. Specieller Teil. Jena. Beiheft: 1-60. pis. Mobius. 14 :x. 1922. 160. J.." Jahrb. Bull Museum Comp. 47. 1908. iii-xiv. 1895. 20 pis. 1912. "Die Bevolkerung des Ozeans mit Plankton. Ernst. Ortmann..: 1-120. Macmillan: xx. Fischer: vi. W.. idem. 22:16-109. 332 ff. E. 726. 1893.. 10:601-682. viii. Idem.298 13... Hans. Plankton-studien. 7 pis.. The depths of the ocean: a general account of the modern science of oceanography. Ernst. A. Beiheft: 1-26. 1896. L. W. Wiss. 221 figs. (pt. Aulacanthidae-Concharidae. Jena. zool France. deut. "Beitrage zur Kenntnis der chemischen Zusammensetzung wirbelloser Tiere." 105. 3 text figs.. Richard. Komm. .. 22." Ber.: 1-148. Flarnmarion: iii. 3 maps (p. Tiefsee Exped. and 656). fishes of N. Anst. 61). 6 figs. "Ober den Bewuchs auf den treibenden Tangen der Hamburg." in Beitrdge zur Kenntnis des Meeres und seiner Be- Lo wohner. pt. H.). 1 map (p. 1914. New England. "Crustacea Malacostraca." Ergeb. Exped. 1890. pts. 1873. Haecker. 1921.. H. 6:322-324. "Die Appcndicularicn der Plankton-Expedition. 2.. Ergeb.. 336. 19. 75). "The North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America north of the Isthmus of Panama/* Bull U." Naturwissenschaf- ten. Hamburg Wiss. 2279). Gran. 3 maps (p. The Pelagial Fewkes. p. 28.). "Zen trifugen plankton und Hochseestromung. Jordan and Evermann. London. 4. 3313. Karl. 5 4 figs. 821. 1904. 2 pis. (pp. H. 25. 1 text und Schizopoden. 24. 23:98-104. Hentschel." Verhandl Lohmann. Ret. Hyman. pt. Y." 29. Plankton fig. 115).) 16:233-282. 13 text figs. 1912. Valentin." Butt. 16 pis. 2.. 26. 1920. Intern. zu Stockholm. 1919. 612 ff. "Neu und altbekannte Ascidien aus dem Reichsmuseum Jahrb. "Die Wirbellosen Thiere der Ostsee.. and B. 21. 1. "t)ber Schwimmen der Fisehe. 3." Arch. illus. Idem. Helland-Hansen. 3. 1918. 62 pis.. Joubin. "Note sur divers fragments d'un ccphalopode: Attoposis mollis Vcrrill. 2:1-120." Wiss. 391 pis. zool Ges. Meyer.. Bianco. (p. Idem. Erste Liefcr- ung. Louis. soc. 1940. 16. Heft Gb. Anst.." pis.. "Die Probleme der modernen Planktonforsrhung. 1908. S. das 17. 13:209-240. 18. A. 1-4: 20. 15. H. illus. Kielcr 30." Hydrobiol. f 38. Meeresunters. 1897. Appelof. ilia* (p. Salvatore. Natl Museum. Murray and Hjort.. vol. illus. Cephalopodes du musee poly334. pis. McGraw-Hill: xii. 261). 14. sur divers cephalopodes. With contributions from A. 91.f. Sargassosee." Plankton Ergeb.. 23. Hesse. (p. Exped. "Dekapoden 31. technique de Moseou. . "Pelagische Tiefseefischerei der Maja in der Umgebung von Capri. Ix.

. illus. 1896. 339).. E. (n. 5:1-136. 1 map.). "Function of spines of crustacean zooca.. Kristineberg och 4... Beitrage zur Morphologie. [Quoted in Murphy.: 1-476. 7:345-394. "Om utvecklingen af Sveriges zoologiska hafsstation 39. A. 42. 21)... Herkunft. Leipzig. 2 pis. W.. 19:523-526. 2. Steuer. Plankton Exped. 18 figs. J. 1878. Zoo/. 1 pi. 14-17. 36. Weldon. The Atlantic: a preliminary account of the general results of the exploring voyage of //. 1910. 2 vols. 41. 723. Planktonkunde." Zoologica sachen der sogen.M. 1906. 39 pis.) 1:169-170. 2.." Ibid. "Die Acephalen der Plankton-Expedition. 5 pis. 35. F." Zool Anz.. N. Harper: 2 vols. Challenger during the year 1873 and the early part of the year 1876. 11 figs. L. F. ton-Expedition. illus. (Stuttgart). 44. 1890. 1917. Theel.. Protistenk. Okeanographia. 1896... "Noctiluca miliaris Suriray. (p. 43.: 1-44. naturforsch. Woltercck. "Schwarmbildung in Meere.: 1-33.. 1936. Ges. 1921. American Museum (Chapter 23)]. d. Teubner: xv. L.." Marine Biol Assoc. 5 pis. Morphologie und Physiologic ( Beobachtungen an der lebenden Zelle). 1908.. N.. United Kingdom. Hjalmar. 34. Rudolf von. 33. "Die Foraminiferen ( Thalassophoren ) der Planksiologie. Rhumbler. W. Petrograd: 2 vols. A. Heft He.. of Natural History: Wege biologischer Mittelmeerforschung. "Ueber Funkion. Artzte. "Die Chaetognathen der Plankton-Expedition. 37." und Cytologie: I." Arkiv. pis. Adolf. Ritter-Zahony." text figs. Simroth. Jules. 1911. Heft Fe. No. Idem. Oceanic birds of South America. illus. 26:474-548.S. Popofsky.s. 3. 1 Ergeb. 41 figs. 365 text figs... (p. 40. 42:1-98. Y. Schokalsky. 9.Bibliography 32. 299 "Ubcr Acanthometriden des indischen und atlantischen Arch. Heft Lc. Y. 85:170-198.." Ergeb. . (vol. 1911-1913. C. R. PhyOceans. Vanhoffen. pp. 2 maps (p. und Entstehungsur'Schwebe-Fortsiitze' pelagischer Cladoceren." Ibid." Verhandl. 1913. Pratje.. Plankton Exped. 38. Thomson. R. "Ziele und om djurlifvet i angriinsende haf och fjordar. Heinrich. 1913. 175 map 217 ff. 2.. 41).

The penetration of light directly governs the presence of plants. The lighted stratum is commonly divided with the latitude for different localities. 185). in which even the great increase in pressure seems into a euphotic stage (0-80 It to have little effect on distribution. for to greater depths in the tropical Lohmann's figures. Any m.4 plants. In both the benthal and the pelagial they are principally developed in the upper stage of the lighted stratum. and extend than in the polar oceans. primarily composed of Average Numbers of Single Organisms per Liler Depth 300 . It is important to note that it is the threshold values for optic sensation and for plant assimilation that are important for animal distribution rather than the minute amounts of light present at great depths.14.) and a dysphotic stage (80-600 m. ami The Abyssal Benthic Pelagic of the Communities Sea THE DEPTH OF PENETRATION OF LIGHT INTO SEA WATER HAS BEEN DIS- cussed previously (p.). locality varies in depth diurnally 16 The lighted stratum at a given also varies notably and annually and sharp delimitation is made still more impossible by the varying threshold values for different animals and plants. does not seem practical to attempt a further subdivision of the lightless lower stratum. are as follows: nannoplankton.

p. however. (i. centages correspond to various depths: 15.5% 5460-71280 m. 400 m. but divides into pelagic and benthic same way as the lighted stratum. say.. the subantarctic intermediate The whole water at depths of 1000 to 2000 m. A great number of animals directly dependent on living chlorophyll-bearing plants for their food are thus confined to the euphotic stage. -"' More species of animals were secured in the uppermost 180 m. 0-180iii. are poor in life.34% 1840 -3640m.). in a vertical distance of 3000 m. 44 fishes and 35 shrimps were The Foraminifera collected by the Gazelle 12 were as fol- Depth in mHers of species 100 138 100 500 500-1000 134 1000-2000 147 Number Depth in 354 meters of species 40003000 53 30004000 79 4000-5000 38 5000-6000 19 Number The great depths of more than 4500 m. 6. in the benthal as well as in the pelagial. Beebe's 3 direct observations down to 924 m. to 1500 m.59% Depths of H). Nevertheless. 8 The animal life of this vast area. decreasing with depth. in contrast.42% 3640-5460 m. constant low temperature. exhibits a much greater uniformity of environment than the much smaller lighted stratum absence of sunlight. 32). per day. a number of special characters are common to abyssal animals of both divisions. Both number of specimens and number of species regularly de- crease with depth. region below. net hauls of the Michael Sars produced 10 fishes and 11 shrimps from 4500 m. and the most successful dredge-hauls secure only a few specimens. this is 2 clearly shown in the total collection of the Challenger expedition.e. in the rest of the ocean together in the secured. indicate that there is a greater amount of fish and large invertebrate life at this depth than is demonstrated . than The vertical (see table. lows: 900 m.. in the is by no means uniform. moves north near Bermuda at the slow rate of 400 to 500 m. from 1350 to 450 m.Abyssal Benthal and Pelagial 301 Greater depths are reached only by the protected spores and dead remains of plants. and almost complete absence of motion. The amount of water in the abyssal stratum exceeds that in the Of the total surface of the oceans the following perlighted one. 58.

(see p. . 192). and the shrimp Gennadas parvus. Pelagic deposits are built up more slowly and are red.302 Abyssal Benthal and Pelagial catches. Such materials are in general deposited near land. the snail Natica groenlandica. may be men- tioned. the bivalve mollusk Scrobicularia longicalranges from lus ranges from 36 to 4400 m. The abyssal benthic community The sunless benthal exhibits different habitats corresponding to the different biotopes of the littoral. their nature varies tive rapidity ental shelf. of varied origin and composition. Even the largest such material beyond the border of the continental shelf at a depth of 200 m. by net His findings. in which case they may be referred to as eurybathic as contrasted with steno1S that descend below bathic forms. greenish. deposits The sea-bottom that Deposits beneath the open ocean contrast with terrigenous ones in many are composed of the calcareous or siliceous skeletons of The with locality and with depth. 12 also occur in the upper 180 m.. yellow. fish Pyrosoma elongatum between 750 and 1500 m. or gray in color. The ascidians Caesira eugyroides from 450 to to 540 m. Examples of stenobathic forms.. from 4 2300 m. the former derived from land areas and made up of decomposed rock. but they may occasionally be carried to great distances by iceisolated places bergs. Rock bottom is present in deep water only at very where special conditions cause a current at depths of more than 400 m. from 600 to 5600 m. do not alter the conclusions just given. and the Argyropclecns hcmigyvnnus from 150 to 500 m. With these exceptions the bottom of the sea is covered with deposited materials. principally based on differences in the substratum. The coarser deposits are laid down closer to the land. various types of bottom deposits intergrade with one another. are few. Thus of 20 chaetopod worms 1800 m.. though not in sufficient quantity to produce a uniform deposit. those based on marine organisms do not predominate in the continpelagic organisms. and abyssal terrivers carry little of rigenous deposits consist only of the finest materials. principally quartz sand. brown.... Animals are frequently not confined to any given depth. Echinocardium autfrale to 4900 in. Murray distinguishes terrigenous and pelagic deposits. aside from purely littoral or purely surface animals.. blackish. however. Deposits of terrigenous origin are built up with comparaand tend to be bluish.

Globigerina ooze. facilitated. crinoid sea. This fact may be in part due to the great depths in which the clay is found. broken only by isolated areas of red clay. where diatom skeletons composed the principal deposit. In the East Indian waters. FIG. diatomaceous . Chun reports that the river of the Gulf of The may mud Guinea is poor in animal life. and second only to red clay in extent. covering 29.2% of the ocean bottom (105. Globigerina ooze is widespread. as in the East Indies. on account of its lime conis especially favorable to the development of animal life. poorly developed. It prevails especially in the North Atlantic. The calcareous deposits are developed especially in subtropical and tropical where lime deposition by animals is regions. 65. The glass sponges (Hexactinellidae) 29 with their siliceous skeleton do best on siliceous deposits. Single groups may thrive on a special type of bottom. At very great depths the pressure prevents the deposition of either lime or silica. which are redissolved. which is the typical is deposit of the Pacific. Rhizocrinus a the stalked lofotensis. km. tent.The Sea-bottom Deposits or white. Thus among the echino- derms there are some very thin-shelled and irregular sea urchins and an excess of holo3* Life on the red clay is everywhere thurians. and the only deposit is a red clay. consisting of shell-less holothurians and worms. of deep After Boas.000. tively shallow Pteropod ooze is found in relawater and relatively small areas. The siliceous deposits appear in polar regions and in regions where an abundant influx of argillaceous material supplies the plankton with silica. At certain of the Challengers stations in the Indian Ocean. they have a rich fauna.). and diatomaceous and radiolarian ooze as siliceous. the product of the decomposition of volcanic materials. 33 303 Murray distinguishes pteropod ooze and Globigerina ooze as calcareous. On the contrary. terrigenous deposits vary in nature. where such deposits are abundant. the higher animals contained only small amounts of lime in their shells.000 sq.

304 Abyssal Benthal and Pelagial '". sea. Chlidonophora chuni. Eurycopenovae-zelandiae from 2000-m. 67. Deep-sea isopod blue ooze. FIG. depth. I FIG. 66. . on After Beddard. a long stalked bruchiopod from the deep After Blochmann.

pelagial varies from 200 to 600 m. a quartzitic terrigenous deposit. 68. butter in summer. 67-68). Deep-sea crab Phitymam icyviUc-thomsoni from 275-800 m. appearance and consistency to require some means of support. No sharp faunal division at this the inhabitants of the abyssal point is possible. and blue. third. 66). the terminal joints of which are expanded by hairs to increase their supporting surface (Figs.The Abyssal Pelagic Community ooze ranking the last is 305 first. especially as many of waters rise to the surface at night. one-third natural size.. a distribution of the weight to widely separated points. After Doilein. radiolarian ooze next. and certain crinoids (Fig. Almost deep-sea sponges are stalked or provided at the base with brushes or collars of spines/ 15 The root-like growths at the base of the all stems of stalked forms serve the same purpose. The abyssal pelagic community The upper limit of the sunless in depth according to the latitude. and. deep-sea ooze. horn corals such as Chrysogorgia. are associated adaptations. 65) and brachiopods (Fig. in animals with legs. As compared with their relatives elsewhere. as in the hydroid polyp Branchiocerfanlhus. FIG. An enlargement of surface. while others undergo their develop- . in Deep-sea ooze may be compared Animals living on it Thus flattened forms such as abyssal sea urchins. crustaceans living on the ooze usually have very long limbs. Sperosoma grimaldi are found among Holothurians have a specially broadened sole.

The food of the deep-sea forms. an archaic cephalopod. Nocturnal net hauls on the surface frequently obtain fishes with every mark of the inhabitant of abyssal waters. in 20 seconds. 31 common to pelagial and bcnthal In spite of the interdigitation of the deep-sea fauna with that of the lighted zone. from 6 to 8 P. 1 * During the day.. Animals of deep water may come to the surface for egg-laying. the uniformity of these factors being more marked than in the rest stillness of of the ocean. it is taken at 350.M. Living plants are not available to deep-sea animals as food. only scavengers and predators are represented. about midnight the whole population accumulates between 46 and 3 m. its abundance depending on the number of animals rate of sinking of the food materials is slow. depth. an inhabitant of great deeps. The low in the upper zones. Cod caught on hooks at depths of 360 m. is derived from the bodies of the surface plants and animals that sink to the bottom. In the herring fishery. is such an exception.. The details have been ascertained for Calanus finmarchicus. the majority are at a depth of 180 m. and on the falling excrement of the surface animals.M. to 6 P. to the surface. salpa of 5-cm. whose larvae are taken at depths of 1000 m. and the great pressure. the the water. It is unusual for a surface animal to develop in the depths. Food falls like gentle rain into the depths. Herbivores are consequently wanting in the deep-sea fauna proper. the absence of light. by day feed on cephalopods (Qmmastrephes) which are taken at the surface at night.306 Abyssal Benthal and Pelagial at the surface. The A . the peculiarities of the environment to which all deep-sea creatures are subject result in many parallel adaptations.to 450-m. ment There are forms that remain permanently in the abyssal waters. the low temperature. length sinks 40 cm. the bottom trawl is used by day and surface nets at night.M. except as they prey upon each other. thus whereas 90$ of the fish larvae and juvenile fishes collected by the Michael Sars in the Atlantic were caught in depths of 0-150 m.. The siphonophore Velella. from 6 A. The composition of the surface plankton in consequence varies from night to day.M. The peculiarities consist in the nature of the food. breeds in shallow water near Amboina from Abyssal characters May to September. Alepocephalus appears to develop entirely below the lighted level. at which rate it would require 2 days and 7 hours to reach a depth of 4000 m. and between 4 and 6 A. Smaller animals also make these periodic ascents and descents. it is uniformly distributed from 350 m. Nautilus.

and cirripedes live on the detritus. Predaceous deep-sea forms require characterized tion. example. Saccopharynx. lamellibranchs. Hydroid polyps. worms. the shark Scapanorhynchus. the more con- spicuous. Lamellibranchs and snails are often minute. a meter in length. so that their evolutionary transition to the abyssal benthal 26 was simple. is large. holothurians are the most abundant deep-sea animals. is an exception. The Pleuroto- midae have given up their predaceous habits in the deep sea/' Amphipods and isopods are already detritus feeders and flourish in the deep sea. sponges. feeding elongated. Deep-sea animals are usually small in comparison with their relatives at the surface. Chimaera. than is to be found in the depths beneath the 2 open ocean. Fishes are relatively small. their little comment. Eupharynx). on the botforms the food supply of the benthic animals of the deep Numerous species of Foraminifera live on it.Characters Common to Pelagial and Benthal 307 temperatures of the deeper water prevent decomposition. is in comparison the size of their relatives in shallow water. produces a greater supply of food in the littoral depths. the less of this food supply reaches the bottom. which is collected by self-made water currents. even in this time. In general the predaceous animals of the depths display few special characters and no special superiority on account of the greater severity of competition. their radula reduced. long. The remnant of the falling food material accumulates tom. This must be one of the reasons for the reduction of popu- way by lation density with depth. several meters The reason may lie in the scarcity of food. Some fishes are and wide mouths very strong dentiby extraordinarily distensible stomachs sometimes containing prey larger than themselves (Meltmocctus. and these may bear special lights. Many echinoit derms have the same feeding habit in the lighted zone. The greater abundance of neritic than of pelagic animals. plus additions brought in by streams and winds from land. The falling food supply is continually diminished on its various anitnals of the intermediate zones. Their digestive tract is habits. There is a notable increase in the number of structures that are inter- preted as lures. rarely reach moderate The with relatively gigantic size of a few deep-sea forms. where sea. and size. Many snails adopt similar for the Trochidae. and hence a greater concentration of animals. and the excretory tube is elongated and carries the excreta away from the feeding ground. Long duration of growth correlated with postponement of . so that under equal conditions the deeper the sea.

for pelagic animals of diverse groups give off light in sufficient quantity to produce the bioluminescence of the surface of the sea. The density of the animal population on or near the bottom in deeper water is unknown. depth off Bermuda. positive correlation be- tween the amount of life at these depths and the development of ultimate source of food for these utilization of dissolved substances.35 gm.308 Abyssal Benthal and Pelagial sexual maturity. at least to some extent. The lights can be concealed or opened to flash like so many fireflies. m. These quantities are approximately 300 times the average amounts in the dry substance of all the marine organisms present on the average in a cubic meter of water. The deep-dwelling animals appears to be from (a) surface organisms or which sink. with Pyrocystis in the tropics. their excreta. The small number of bacteria in sea water argues against their having a high food value. These microorganisms. but the excreta and bodies of large animals must provide a considerable amount of food. There is a definite. the jellyfishes Cyanea and Pelagia. the ctenophores Beroe and Cestus. per cu. together with the particles falling from upper levels. The theory of Putter that organisms take such dissolved nutriment directly from the water may hold for smaller forms such as bacteria and for some small protozoans. Krogh found that the concentration of nitrogen and carbon was constant from surface to bottom at 0. 20 Bioluminescence The production of light by deep-sea animals is to be regarded as an adaptation to the absence of sunlight.. In a column of water taken at 5400-m. or ( b ) the Plankton organisms cannot reach the bottom at great depth in significant amounts. copepods among crustaceans. The protozoan Noctiluca. and to feed upon other minute particles. Some shallow-water fishes have light-producing organs in which luminescent bacteria are contained in a palisade of tiny tubules. Light production is by no means confined to the animals of the deep sea. Calculations of the number of whales dying in Antarctic waters support this contention. respectively.244 and 2. has been suggested as an explanation. Phyllirhoe among snails. those in the bottom ooze may be present in high enough numbers to have significance as food for protozoans that are also able to utilize dissolved substances. phytoplankton at the surface. probably constitute the base of the food pyramid in the deep waters. . and Pyrosoma among tunicates are examples. conditioned in turn by low temperature.

littoral benthos is exceplight tional (Pholas. the others appearing momentarily general color. these sent forth flashes of light so bright that the 20 torches lighting the work paled into insiglike "How nificance as soon as the polyps were near them. to be a social signal in the pelagic schooling Animals that live in the dark Anomalops. which give lights at For the most part. off light-producing mucus from their among the macrurids. . 924 m. a dipterous larva (see Chapter 27). whose intensity then increased. occupying the twilight region of the sea. Bioluminescence is more common among crustaceans. from blue to various tones of green. and Gorgonidae. All these have usually a continuous light.Bioluminescence 309 1 ** They seem production. whose clarity and beauty of color delight every observer. including an earthworm and the various light-producing insects. and sometimes to the white of flowing iron. De Folin 15tt expresses his impressions of nocturnal dredging as follows: great was our astonishment when a great number of bouquetgorgonids were taken from the net. than in the lighted surface zone above or the bathypelagic No less than 44 per cent of the fishes of depths 27 It is estimated that two-thirds m. Beebe found the first animal 30 region below. and cephalopods of the mesopelagic zone. although there are a number of bioluminescent forms among terrestrial animals. The largest number is perhaps supplied by alcyonarian coelenterates Alcyonidae. may show no tendency toward light Cave animals exhibit only a single known light-pro- ducing form. Meanwhile the predominant color was plainly less. and annelids are represented by Chaetopterus and Polynoe. passing from violet to purple. from red to orange. became green. light production may nevertheless be said to be especially characteristic of of light deep-sea animals. body. special In some forms these organs are simple in structure and are mere acskin glands. Pennatulidae. however. and thereafter in slowly increasing numbers down to Diffuse luminescence occurs the greatest depth reached. 207 m. The production by animals of the . 900 are light producers. Echino- derms have the starfish Brisinga and Freyella. By direct observations." In all these animals the not confined to special organs. but is and melting quickly into the power of light production is more or less diffused over the fishes. Chaetopterus) the abyssal benthos swarms with producers. beyond of all bathypelagic fish species and over 96 per cent of the individuals 3 are luminous. From every point of the main stems and branches beams of light radiated. the pelagic forms have luminescent organs under the control of the nervous system.

fibrillous in c. in c. 69. lac. and c. of a fish (Argyropelecus affinis). ~ lacunae. b. schizopod crustacean ( Nematoscelis mantis ) . reflector lined.310 Abyssal Benthal and Pelagial Lens Light organs of deep-sea animals: a. of an octopod ( Abraliopsis morisii). Light cells stippled. backed by a pigmented layer. b and c after Chun. a after Brauer. with an afferent nerve. A lens of varying structure in front of the light cells. . of a black. n. made up of cells in a t homogeneous in b. in b. FIG.

In other more complex and reaches its highest de- among Euphausidae (Crustacea). this light is weak at of alcyonarians.Eyes cumulations of gland gree in Relation to Depth 311 cells groups the structure becomes with a luminescent secretion. covered over by a pigtapetum. In pelagic animals. Oases of light occur where there are forests of gorgonians or meadows The In comparison with sunlight. the highly developed light organs must have other uses. 33 of bottom-dwelling fishes may be well own light-producing organs are lacking. but it does enable animals to direct their motions by means of their eyes. Behind the group of light-producing cells is located a in the form of a concave reflector. Many luminescent cephalopods and fishes maintain themselves in the transition zone between the lighted and the abyssal pelagial. This helps explain the great variety in arrangement of the light organs and in the color of the light emitted. and of the species in gregarious forms. Eyes in relation to depth Oddly enough. cephalopods. fact that the degeneration and loss of eyes is less general in explains the abyssal fauna than in caves. 69). What may be the function of this bioluminescence? The attraction of prey must be considered as the primary possibility in the benthic forms. best and does not reach far into the water. formed in different ways in the Independent convergent origin of light organs may be seen groups of cephalopods and fishes. the eyes developed even though their Apparently these brates. Many are grouped around the mouth. and bony fishes. ment layer. but scarcely touches the problem of bioluminescence of alcyonarians and other sessile forms. Convergence in the evolution of these organs in such diverse groups is notable. It is probably equally important that their characteristic arrangement makes possible a recognition of the sexes. and again suggest a use as food lures. shallow water. off fishes use the light given by benthic inverte- . It is notable that light production is unknown among the cephalopods and wi fhin fishes of different animals (Fig. Some of the relation- ships are indirect. Thousands upon thousands of luminescent animals provide torches that light the abyssal depths. An outer lens is present. which are absent in the few coastal forms. This light is not uniformly distributed. thus a hermit crab is known that carries luminescent 2 The development of bioluminescence actinians on its borrowed shell. sunless depths of the sea are consequently not without light. The deep-sea members of the Iniomi (lantern fishes) have luminescent organs.

Brama Aphanopus carbo. These are found only in the smaller fishes. 72). 0. Pterygioteuthis. schizopods (Fig. Enlargement of the lens alone. whereas the lateral portion has shorter and strongly divergent elements. cephalopods and FIG. 71fo). Gigantura chuni (Fig. much zones. depth. exhibited by the so-called telescopic eyes (Fig.312 Abyssal Benthal and Pelagial of The eyes 70a). Macrurus fasciatus from b. such as geckoes. The deep-sea . and sergestids. and tarsiers. but their number in the pelagic formation is small. Such eyes are otherwise unknown in these groups. 73&). A number of 2 decapod crustaceans exhibit this peculiarity (Fig. 5 36 * There are eyeless animals in the deep sea. abyssal also have en- larged eyes. many mesopelagic animals are especially large (Fig. larger than in their relatives of the upper or lower depth These animals live near the upper limit of the sunless depths and thus correspond to the nocturnal or twilight terrestrial animals with large eyes. depth. Their eyes are divided into a dorsal portion with upward-directed elon- gate and little divergent facets. Eyeless animals and those with reduced eyes are numerous in the abyssal benthos. that this in adaptation to meso- pelagic life. 7la). 250-in. affords a second type of adaptation to the dim light. the retina of the deep-sea fishes of all groups has specialized light-sensitive rods. longipinnis. Bathyonus tacnia After Giinther.8 cm. As a further adaptation to the small amount of fishes. 70. for example. and Certain mesopelagic like Chiroteuthis Regalecus. but their habits are insufficiently known is to make it certain from 4500-m. Such telescopic eyes (really tubular) have developed independently in five different orders and eight suborders of 2 7 21 The deep-sea cephalopod also has eyes of this type. have very large eyes. the cones are largely or wholly absent. similarity of the frontal eyes of pelagic deep-sea crustaceans is only apparent. of which the largest. A ' - abyssal light. as do many amphipods (Fig. owls. although there is also a relative enlargement of the lightgathering surface. Numerous fishes of the border zone of the. measures 11. instead of enlargement of the whole eye. 71d).

Pandalus magnoculns.Eyes in Relation to Depth 313 FIG. xpinosuni. Nr. b. c. C. Cyrtisoma Deep-sea crustaceans with enlarged eyes.. in which F is the anterior and S the lateral eye. d. lens. cornea. /. Telescopic eye of the deep-sea fish Argyropelecus in transverse section. u-. a hypcrine. After Brauer.stigophorum. report. . retina. a caridid. with the dotted outline. Thysanoessa gregariu. 72. an cuphausid. c after Chun. eye of an euphausid. Stylucheiron ma. . L. in vertical section. a. [/. and d from the Cliallenger 71. R. pigmentary layer. \\ R y Fic. of a normal eye for comparison.

Argyropelecus Gigantura chuni. In correlation with the difficulty of orientation by sight in the dim numerous deep-sea crustaceans and fishes have antennae of unusual length in comparison with those of their relatives in the lighted zone. Fishes and cephalopods rarely exhibit complete degenera- tion of the eyes. Winteria telescopa. Those with large eyes live in the twilight zone. reduction of the eyes with increasing depth is found within the same FIG. c. Among fishes the Macrurus species of the Challenger collection may be sorted as to depth according to the size of their eyes. has antennae four times the length of the body. known from An eyeless cephalopod.314 Abyssal Bcnthal and Pelagial The eyes of benthic pectens and Eulima and Fusiis are eyeless. Munnopsis longicornis (600-800 m. is a depth of 1500 m. After Braucr. b. in of facets and in both number crustaceans are frequently reduced. times the length of the body. Sergestes magnificus (from 800 to 1200 m. n found all crabs from consideramount of pigment present. 73. Doflein able depths with fewer facets in their eyes than the related littoral forms. the antennae are more than eight times the body length. Telescopic eyes of deep-sea fishes of various families: a. and in the shrimp Aristacus they measure .) has antennae three light. Arachnomysis. depth). Cirrothauma murrayi. affinis. and 20 per cent of the higher Crustacea in the catch of the Among some crabs Investigator had pigmentless or reduced eyes. in the isopod. with well-developed eyes. 2 species. those with smaller eyes in more considerable depths.

In the . besides dark brown. single rays of the various fins serve as feelers and are longer than the body. with dim light. and only the silvery gleams from their sides make them visible. After Gunthcr. dark violet. on the The edge of the disk of the medusa Rhizostoma. 74). mackerels. makes red appear as black.Coloration in Relation to Illumination 315 10 to 12 times the length of the body. There is a decided predominance of red colorations in all tones. Coloration in relation to illumination 2fl protective colorations of the animals of the lighted benthal region have been discussed previously. For the rest. Similar develop(cf. depth in the soutli Pacific. the fishes are predominantly silvery. In the lower stages of the lighted zone. Bathyptcrois Umgicatula from 4HOO-m. in correlation with the variety of background on which the animals live. 53a). colorations of pelagic animals in the lighted zone are much The plankton animals are in the main transparent. where sunlight is absent and where a uniform ooze covers the bottom. The dorsal half of pelagic juvenile fishes (Mullus) and of adult fishes like sardines. In greater depths. the decapod Virbius. They are extremely varied. ment of the organs of touch and smell found in cave animals Chapter 27). and in the snail Glaucus (Fig. The absence of red light in depths even of 50 m. In numerous fishes. part owing blue is the dominant color. in the copepod Anomaloccra. such as Bathypterois (Fig. The less varied. and the flying fishes. The dark brown back of the herring is invisible from the surface. and black. colorations are much more uniform. as in the siphonophore Velella. line system is also notable in many deep-sea is fishes. High development of the lateral FIG. 74. is blue. as shown by Fol in diving experi- ments in the Mediterranean. Some Foraminifera and black striking for their dark reddish violet of the deep sea are coloration. in to the high water content of their bodies.

The same colorations are found in The scyphomedusan Atolla has increasingly the lightless pelagial. There are also. are characteristic of the abyssal waters. The deep- sea nemertean (Pelagonemertes). and the fish Bathypterote longicauda (Fig. and 200 m. The wonderfully fragile glass sponges (Hexactinellidae). the bony fishes. the annelid Tomopteris (Fig. and red is a rare color among them. The colorless or the surface. slender. Macruridae (Fig. and long-legged crabs like Kaempfferia require quiet water. 2 ' 7 ' 21 cotta in color. green. The starfish of the deep sea are red. in the tropics. 500 m. whose lower limit coincides with the upward range of the former species. the water has motion. signata. colorless thuria (Fig. Delicate. in the temperate zones. and red crustaceans are shells of snails and lamellibranchs. is light-colored. 75). are likewise characteristic of the depths. the long-stemmed crinoids. The drop low temperadifficulty tures in the great depths of the abyssal ocean. and brown forms the littoral. in polar seas are uniformly red. and a number of pelagic cephalopods are red. in correlation with their movement and fragile also weak powers of locomotion. to be sure. from deep water. long-stalked animals or animals with an ungainly walk on stilt-like legs are not uncommon. Cyclothone microdon. is black. The absence of water makes possible the survival of many forms with weak skeletons. dark coloration with depth. while C. and the band-like fishes Regalecns and Trachypterus. the deep-sea pelagic holothurian Pelago- deep rose. Gigantura. some arrowworms.*and Gastrostomidae. and intensely red of alcyonarians. 53c). 76). decapod crustaceans below 750 m. 74). or terra Red cephalopods are not rare. whose presence is associated with the difficulty of temperature with depth leads to uniform of lime formation. however. are mostly have pale coloration. such as Chimaera (Fig. Chlamydoselachus. The bathypelagic larvae of Vclella are red. 59) all is Almost animals in the abyssal pelagial such as the crustacean Scrgestes inagnificus. such as the gigantic Branchiocerianthus imperator. The in . Compressed forms such as the deep-sea shark.316 Abyssal Benthal and Pelagial abyssal benthal are red hydroid polyps. Fishes with tails drawn out into long points. bright red sea anemones. in contrast with its glass-like relatives of abundant. Body form and skeleton little In depths below 100 m. orange. in contrast with the yellow. Many deep-sea copepods are dark violet. This makes possible structural forms impossible in rapidly or violently moving water. The abyssal fishes are mostly dark violet or black.

Pelagoihuria. 207) is thus repeated in the deep sea.. After Boas. is identical with S. Numerous echinoderms. 75. Such forms all FIG. mostly below 1250 m. polywith a reduced shell. Even in the same reduced in deep water. Accordingly. the starfish Brisinga has a reduced skeleton.Body Form and Skeleton 317 the formation of lime produced by the low temperatures in the polar seas (see p. 300 m. ani- mals belonging to groups with a strong calcareous skeleton or shell in the warmer waters are found with weakly or completely non-calcareous skeletons in the deep sea.. 18 Mollusk shells 400 below from m. the skeleton may be. normal shell. Scalpellum species. Macrurus rupestris. Chimaera monstrosa. After Hentschel.. with a . and the pelagic deep-sea holothurian. Many deep-sea fishes exhibit stearnsi in 200-450 m. have a leathery non-calcareous skeleton. Thus calcareous sponges are wanting below Fir. belong to great depths. 76. has no calcium bodies in its skin. such as the echinothurid sea urchins. morphum from great depths are mostly fragile. Numerous deep-sea barnacles of the genus Scalpellwn have an incompletely calcified shell.

they followed at steadily increasing depths on both sides of the ocean. and Naticidae. abyssal in warm seas. The bottom is smooth. Lithodes.f and of anomuransj inhabit the deep sea in low latitudes. At great depths the water is uniformly cold. where they are absent at the surface. the difference between littoral and abyssal animal life is greater in the tropics than in arctic seas. Pasiphaea.. without With holes or cracks. however. with little motion.* of crabs. Lima. as in Chimaera. This is true of many snails and bivalves of the may be northern Atlantic. The lamellibranch genera Yoldia. the deep-sea fauna is characterized by great uniformity over extremely wide areas. the bottom covered with ooze whose main variation is chemical and which is uniform over wide areas even in this respect. Latreillia. Some deepon the sea forms.318 uncalcified or Abyssal Benthal and Pelagial weakly calcified skeletons. Helena (to 2000 m. ). Nucula. have at least one or a few species near the surface in the Arctic Ocean. .) and to the West Indies and Pernambuco (to 800 m. living in the Arctic littoral to a depth of 50 m. do not enter polar surface waters. for the most part abyssal fishes are archaic. affinities of abyssal Faunal animals The occurrence of surface forms of polar seas in deep waters in the warmer zones. 2 17 ' the Arctic. contrary. The deep-sea genera of shrimps. Centroscyllium 14 is also found nearer the surface in polar waters than in the tropics. without sunlight. The sharks have similar distributions. 4 On the whole. Among many animals the connection between the polar surface forms and the tropical abyssal ones can be followed continuously. to the Canaries and St. especially among the Squalacea. 5G Uniformity in abyssal waters Uniformity of environmental conditions in the deep sea is greater than in any other division of the ocean. Crangon. which also has characteristic northern snails of the families Pleurotomidae. is explained by the temperature relations. are not deficient in calcium carbonate. Maia. the same is true of decapod typical northern genera of shrimps. environmental barriers wanting. Trochidae. since there are no modeling forces in play. The great majority of deep-sea fishes. and Abra are littoral in crabs. and it was even supposed at one time that it was essentially uniform throughout the * f $ Homola. 23 The northern starfish Brisinga is found in the abyssal Indian Ocean. Pandalus. Hymenodora and Pontophilus.

world-wide deep-sea decapod crustaceans belonging both to the pelagic and benthic Silenia sarsii habitats.. Cyema atrum. in all seas. Of 523 species dredged from depths more than 1800 m. as Chimaera monstrosa or Cyclothone microdon. Melanocetus krechi. is not world-wide. Mastigoteuthis belone. Numerous species also have the same distribution. predominantly the bathypelagic animals that exhibit such wide The distributions. and others. and the great number It is makes any faunal of animals of various groups found in the depths of all oceans division of the abyssal oceans more or less vague. Of 130 bathypelagic genera of fishes a fourth are The flammea. Ortmann 28 enumerates 49 species of Atlantic and Indian oceans. 603? were unknown from other regions. from depths greater than 2300 m. from the Atlantic and Indian oceans. 19 The 21 North Atlantic species of sea urchins and the 28 west African forms have only 10 species in common. south of the tropic of Capricorn. though their genera and families are much the same. only 363? were known from other regions. faunal differences are no demonstrable barriers. On the other hand.Uniformity in Abyssal Waters oceans. known from three oceans and another fourth from two. 319 The increase in our knowledge of deep-sea life has shown that the uniformity. A number of abyssal lamellibranchs such as and Semele profunda have a very wide cephalopods CalHteuthis reversa. Faunistic differences are less marked in the abyssal benthal and pelagial than in the lighted benthal or even in the lighted pelagial. Spirula is known from the Pacific as well. Numerous sea urchins of great depths are common to the Atlantic and Indo10 The deep-sea ostracod Gigantocypris is known from the Pacific. Toxeuma distribution. The east of and west Atlantic in the abyssal zone have a total of 74 species of sea urchins. 26 Although most species of the deep-sea fish Cyclothone are sub- . It is to be watched whether the number of the deep-sea forms known to be widespread will be increased as our knowledge grows concerning the deep sea. but widespread benthic ones are not wanting. and others. but only 24 are common to both areas. The decapod crustaceans exhibit similar differences in the North and subtropical Atlantic. 25 The Sunda Archipelago on one hand and the Gulf of Bengal and Oman on the other exhibit two distinct deepsea faunae of holothurians. and Malacosteus indicus. medusae Atolla and Periphylla are known from all seas. its water masses. are known from the Atlantic and Indian oceans. may appear even when species taken there Of the 272 by the Chal- lenger near Kerguelen. though great. and slow-drifting currents.

is so different from that of the lighted zones in the ocean that there is a recognizable A rapid review of the deep-sea fauna peculiarity in this respect. the temperature is about 6. No Atlantic abyssal species of The genera Cyclothone and fishes are found in the northern ocean. Canary Just south of the ridge are found animals that range to the Islands. the only exchange of forms possible is in the uppermost zone. while in the Norwegian Sea Lycodes (related to the "eel mother" Zoarces) predominates. which play an important role in the Atlantic. a few kilometers to the north occur forms that ex- tend to Spitzbergen and even farther. at an equal depth. of wanting. C. between depths of more than 1000 m. according to the groups of animals represented. and its continuation in the direction of the northeast corner of Scotland. the temperature is +7.-*' Similar differences are also shown in the benthic fauna of the two sides of the ridge. At the deepest point on the ridge. livida occurs only on the west African coast. and . the Wyville-Thomson Ridge. below the surface. at depths 0. North of abundant. Nasselaria and Phaeodaria outnumber the Spumelaria and Acantharia with increasing depth. and although the peculiar adaptations to the deep-sea environment are by no means general. the composition of the deep-sea community.41 prevails. Abyssal animals reviewed Although a considerable number of the deep-sea animals are eurybathic and range upward into the lighted zone. where it is in the deep sea. a temperature of farther to the south. so that a special character of deep-sea life can be asserted only with reservations. This barrier shuts out the whole of the Atlantic deep-sea fauna from the Norwegian Sea. The effect of the latter is the more notable as it exis such a barrier. on each side. Macrurus. Murray records 385 species of animals from both sides Atlantic which only 48 ( 12% ) are common to both the warmer and the colder Norwegian Sea. Among the radiolarians. and scarcely a degree of 1100 m. 16) so that in the Faeroe-Shetland channel.5. tends as a narrow ridge. whose deepest point is 556 m.. are wholly of the barrier. where there are great barriers that are insurmountable for many animals. shows this plainly. so that for stenothermal cold-water forms this ridge is an impassable barrier. The Faeroe Ridge between the Norwegian Sea and the Atlantic. with only a few special forms in the Atlantic. Local specialization accordingly takes place also differentiation is Such further increased this ridge the temperature drops much faster with depth than it does to the south (Fig.320 Abyssal Benthal and Pelagial tropical.

which predominate on the sea bottom on account of their great numbers in the lighted zone. Galatheidae are for the most part deep-sea forms. rare. Of Bryozoa. the chilostomes have than forms fewer numerous abyssal representatives.. glass sponges are almost confined to the lightless Turbellarians appear to be absent. especially imperforate forms with sandy shells. Of 49 genera of holothurians discussed in the Choilenger Reports. Foraminifera are abundant in the abyssal benthal. and among them are only the tubicolous forms that feed on detritus.Abyssal Animals Reviewed 321 the family Challengeridae is confined to the abyssal pelagial. developed. amphipods relatively Stomatopods seem to be wholly absent. Among mollusks. about Lamellibranchs half the species of scaphopods are deep-sea forms. The number of annelids is relatively small. although the brachyuran crabs do not go below 800 m. The Globigerina ooze is composed of the dead shells of the relatively few surface forms. stony corals of and among these the depths. Isopods and with numerous well are species. double the . Numerous deep-sea decapod crustaceans are known. Anornura are not peculiar genera. Certain sea pens (Umbellula) reach great depths. predaceous forms are The Gephyrea of the deep sea are also tube-dwellers such as rare. are few and snails still fewer. while the family Pelagonemertinidae is represented by numerous genera and 6 species in the abyssal pelagial. the family Elasipodidae is confined to the deep sea with the exception of two species. Hydrozoa are scarce in the abyssal. of abyssal cephalopods is relatively large. The most abundant deep-sea animals are echinoderms. they have no characteristic abyssal genera. The abyssal sponge fauna is composed of siliceous sponges. some going to great depths. and 10 are found in both zones. are not rare. 20 are abyssal. Sessile Alcyonaria. Paguridae (hermit crabs) are represented by a number of Pantopoda are poorly represented. 19 are littoral. Phascolosoma. The Challenger Expedition secured only six species of nemerteans from the sea floor. Farciminaria delicatissima to more than 5000 m. like Mopsea and Primnoa. the family Eryonidae is now confined to the deep sea. the benthic the bathypelagic. it has 66 abyssal species. only 6 of the 27 genera being known from the abyssal. The deep water are completely different from the reef corals and are almost always simple. which are mainly abyssal though not wholly absent from the upper zones. Among crustaceans the barnacles are represented primarily by the genera Verruca and Scalpellum. Among coelenterates. to which it is confined. Cyclostomes and cteno- The number stomes are poorly represented.

Among abyssal decapod crabs the representatives of the otherwise Triassic family Eryonidae (Willemoesia. 272 species out of 832 descend below 400 m. Agassiz remarks that l the forms with greatest range in depth are also those with greatest span in time. while 28 of the 91 genera reach depths The number of deep-sea fishes enumerated by Brauer 4 is 309 genera with 1007 species. Echinothuridae. this fauna differs conspicuously from that of the lighted zones. of which 131 genera and 397 species are pelagic. for example) are notable. and Ananchytidae. go deeper than 400 m. 32 ' number 84 Among starfishes. produces the relative poverty of the deep-sea fauna. Among fishes the and the shark related to the Devonian chimaeras. which have 12 The proportion of somewhat smaller. may be named as archaic. the living littoral forms extend only into the late Tertiary. with the exception of ascidians. . stony The sea-urchin families Salenidae. forms with soft rays predominate in the abyssal. though Nautilus may rise to the surface. related to Mesozoic and corals are closely early Tertiary forms. In the groups represented. which reached their maximum development in the Cretaceous. Among the bony fishes. Archaic forms in abyssal waters The necessity for special adaptation for life in the deep sea. The cephalopods Spirula and Nautilus belong to the deep sea.322 Abyssal Benthal and Pelagial of the abyssal species of other families. as they do in fresh water. the brachiopod Lingula. only the two oldest genera. Echinocyamus and Fibularia. Tunicates are few in the deep sea. This is consequently one of the places where ancient types have been able to maintain themselves. the competition between species is diminished. Such archaic forms are not absent from the lighted zoneas illustrated by the horseshoe crab. are now confined to the deep sea. The stalked below 800 m. Stalked crinoids. Chlamydoselachus. of which only a certain proportion of animals are capable. which originated in the Cretaceous. Of the mqre recent Clypeastridae. were thought to be extinct until the dredge brought living representatives to light from the deep sea. as is the great number of such primitive groups as the Peneidae and Caridae. and lophobranchiates and plectognathids are wanting. Cladodus. genera confined to the abyssal. 32 abyssal sea urchins and brittle stars is crinoids are almost wholly confined to the deep sea. A. few spiny-rayed fishes reach the depths. and amphioxus but they are relaMany of the deep-sea tively more abundant in the abyssal depths. which were numerous and widespread in the earlier geological periods. While the struggle with the physical environmental forces is increased.

20 text figs. Brinkmann. 328. 4.. 26 pis. Anatomischer Teil.) 3. Jena. 16 pis. 2 maps. H. 3:xi. 484). littoral belong to more primitive BIBLIOGRAPHY 1." Ticffsec Exued. 9-50. 243.. Fischer: vi. 46 pis. 594. A. 390 text figs. Akad. August. as it does for example to the land fauna in Australia. G. wiss. Tiefsee Exped.. Murray: xxiv. Alexander. Museum 1904. d..). Aus den Tief en des Weltmeers. 46 figs. 15. Beehe. R. 1895. four years with the Royal Indian Marine Survey Ship Investigator. 1906. 1:1-180. Comp. 233. W.. 1934." Ibid. Wiss. 1874 bis 1876 von S. Ludwig. Ibid." Z. II." 6:1-314. 11. Ifg. A. Franz. "Oceanographic work at Bermuda of the Zoological Society. (Ny." pis." Science. August." Miinchen (m.. 1895. . Gazelle [1893].Bibliography 323 their The more recent spiny-rayed way into the depths.. 1902. 16 pis. 1 map. 1874. "Die Tief seefische: 266. 1904. Sch. M. 9. and he evidence gorgonians deep-water as maintains that the gorgonians of the genera. The genus of crabs Ethusa. 1946. pis. "The Punaniic deep-sea Echini. Ernst. R. 112 pis. 58 68 figs. Cooke. 8.. (p. 1906. BraiuT. The deep sea is not isolated. and the way into the depths is continuously available to animals of the lighted zone. Coe. Naturalist. 221). : New York Ergeb. William. 1908. has been regarded an example of a form in process of descent into the abyssal habitat. 2:61-288. 1917. Ehlers. 1:1-432. "Die pelagischen Nertinen ( monographisch darBergens Museum Skr.. London.. Egger. 18:193-458. ZooL.. 2:1- 6. "The means of dispersal of bathypelagic animals in the North and South Atlantic Ocean. aus Meeresgrund-proben gelothet von J.. Ifg. 29 7. 311 10.. "Brachyura. Systematischer Teil. Doflein.ool. Schilderungen von der Deutschen Tiefsee-Expedition. 1-459. 5. Ergeb. figs. the ancient character does not apply to the whole. 1900. Chun. figs.. gcstcllt).. 25:1-102. A naturalist in Indian Seas or." Am. d. 12. 30 In some forms renewed speciation appears to have taken place in the -2 regards the higher development of the abyssal zone. Carl." text figs." Cambridge Natural History." 31:10. "Die Tief seefische I.-n. Alcock.. W. "Die Echinoiden. 15. 2 maps (p. 98 3.. 2. Ifg. 11 figs. in which the same species exhibits reduction of eyes with depth. 7. 13. "Foraminiferen AbhamU. 5. (pp. Dotlerlein. Idem. Kiikenthal as of more recent origin.. 80:495-496. "Molluscs. Agassiz. 238). No. 80:453-460. "Beitriige zur Kenntnis der Vertikalverbreitung der Borstenwiirmer im Meere. Although there is fishes have as yet scarcely found this easily recognized archaic element of the abyssal fauna. Mem.

26. C. Zool. 1912.). 340. Abt. 487). Paris.. 821. "Holothuries abyssales Investigator dans 1'ocean Indien.. Princeton Univ. based largely on the scientific researches of the Norwegian steamer Michael Sars in the North Atlantic." 443. Zool. 6:1-21.M. Schulze. Hclland-Ilanscn. Challenger. August. Idem. Semon. 16 pis. 126:441Arch. Monographs. Harvey. 1913. F. 321. 1896. 5. 1887. "The general conditions of existence and distribution of marine organisms. 16a. 1887. Congr. 1 map (p. E." Biontologie. "Conditions of life at great depths in the ocean.. sammlung: 1913. A." Ann. In the Australian bush and on the coast of the Coral Sea. Ordnungen. Set. John. Acad. Princeton. Richard. faune malacologique des sci. . 41 ff. K. O." Rept. G. 21. "Monographic der Selachier der Munchener StaatsAbhandl. 552. 6th Intern. 1904. 2 pis. Congr. (p. Living light. 24." Naturwissenschaften. C. 221. 4:430-439. 60). 10 pis. pt. oceanog. 1 Monaco . 15a. 1901. 113 text figs. "Report on the Hexactinellidae. Ifg. 328. H. H.-n. I Abyssal Benthal and Pelagial Teil. "Cirripedia pedunculate." Siboga Exped. xv." Abhandl Senck. 1907. fasc. 10 pis. 86 figs. M.." 106 ff.. 4.. 468 ff.S. Zweite Halfte: Malacostraca. Boston: 889-899.). 94 pis. 1934. 1920. Murray. 4:140-151. "Sur 1'aire de dispersion de la grandsfonds de 1'ocean atlantique boreal. Suppl. Grein. 22:xi. John.. 5. Leyden: 99-111 (p and Johann Hjort. Sous les mers. 1889.' Zool. Wiss. Macmillan: xx. R. Ges. 1907. Gran. 45 figs.." Proc. 27. Rev. 4 maps (p." (m. Challenger. 4. R. pt. 19. 7th Intern. "Absorption des Lichts in Seewasser (Erster Teil). London. lenger. Ortmann.)." Intern. 1916. Berne: 610-613. 1901. P. Tiergeographie der Selachier. Krogh. 2. Congr. W. 29. London. Proc. du Talisman. 22.. "The theory of abyssal light. With contributions from A. Hydrobiol." Ecol. 30. 18. Press: Henderson. Esterly.. Sladen... R. 20. Paris. 17. A. illus. "Die Bevolkerung des Ozeans mit Plankton. 1887. 3:1-110.. Engelmann: 1-49. 1895. ChalKept. Hoek.. 1 map (p. E. Compagnes d'explorations du 15. L. Murray..)." J. Sci. Seeliger. Appelof. R. "Crustacea. Munchen Travailleur et 16. 27:xi.324 14. de Folin." 60-62. Vaney. Nutting.. R. A. Willy. (p. 4:657-663. and B. 3:1-617. inst. C. and C. "Report on the Anomura collected by //.. 88 text figs. 28.M. "Report on the Asteroidea collected by //. 259). 1898. "Die geographische Verbreitung mariner Bodentiere. 1283 ff. Engelhardt. 1 pi. Macmillan: xv. 8 figs... Bronns Klassen u." recueillies par 1' C. Tierleben der Tiefsee.. 32. Leipzig. 13 text figs.. Residts Voy. 8 figs. The depths of the ocean: a general account of the modern science of oceanography. illus.S. Lohmann. 25. 31. E. P. 3rd Intern. N. Kiikenthal. Koehler. 21:1-513.). 1899. 31A: 1-127. 1940. C.. 1911. Akad. (p. Hans. C. pi. Locard. "Ergebnisse einer zoologischen Forschungsreise in den Molukken und Borneo im Auftrage der Senckcnbergischen Naturforschenden Gesellschaft. 23. 1 map. Percy.. Results Voy.. Bailliere: xi. "Diurnal migrations of Calanus finmarchicus in the San Diego Region during 1909. O.

The oceans. Y. physics.Bibliography Challenger during the years 1873-1876. ibid. 265 Hjalmar. pt. Hills. G. U. 46 pis. pp. Fleming. 1942. Challenger during the year 1873 of the year 1876.S. C. The Cranbrook Inst.M.... 339-344). Harper: 2 vols. 197 figs. H. 785. Part 2.. 893.. Y. M. and R. 1878. pis. 1942. Science: xiv." Part 1. W.. Thomson. their x. 36. biology. Results Voy. chemistry. Results Voy. Challenger during the years 1873-1876.S. 14:1-290. illus... Sci. Sci... Challenger. "Report on the Holothuroidea dredged by H. Theel. Rept. The Atlantic. a preliminary account of the general results of the exploring and the early part 328 ff. field voyage of H. Johnson. W.M.. H. Challenger. idem. vertebrate eye and its adaptive radiation. and general N. 33. 1 map. 34. 2. figs. 4. Prentice-Hall: 1087. Sverdrup. (vol. Walls. 35. N. 1882. 117 pis. F." 30:xlii. 1-16. 7 charts. 325 Rept. 3:1-176. Bloom- .

which condition the development of recognizable animal communities. by depth. smaller bodies of water such as the Mediterranean. in the final analysis. The distinctions to be made life are extremely simple and consist most plainly in a subdivision of the animal communities of the ocean into those of warm and of cold waters. These are the more distinct when such land-locked seas are connected with the ocean proper only through narrow and shallow straits such as the Straits of Gibraltar. near the mouths of great rivers such as the Amazon or the The geographic distribution of the animal communities of the oceanic pelagial are.D. 326 . presence of fresh water. and in the accumulation of gradation depth. Baltic. very roughly corresponding with the tropical and subtropical areas on the one hand and the cooler waters on the other. determined primarily by temperature. or Black sions can lesser seas. Geographic Divisions of the Animal Communities of UNLIKE the Sea MANY OF THE BIOTIC DIVISIONS OF THE LAND. In littoral areas the relationships are frequently complicated factors. in temperature. type of bottom. these separate the animal communities of the and bays from the oceans proper instead of coinIn these ciding with the geographic divisions between the oceans. they may be compared to the climatic zones of the continents. and similar The major animal communities of the sea are the most extensive of zones. but far surpass them in area and in homogeneity of environmental factors and of associated biota. chemicals in the water. there are distinct peculiarities cially Congo. gulfs. espe- seas. Although physiographic divi- be recognized.I. THE ANIMAL CX)M- munities of the sea are not separated into irregularly placed subdivisions but rather extend as broad zones. especially in with in salt content. Within the oceans similar differences may occur in the littoral regions.

stenothermal tropical life can exist. polar limits of the transition zone.Geographic Divisions of the Sea 327 these vast areas are not determined by and run latitude. according entirely separated. after Joubin. Distribution of reef corals vertically ruled. light broken line after Meisenheimer. Capricorn They are much more closely correlated with The boundaries between the isotherms of the surface waters. Upwelling cold bottom FIG. 77). the basis of the distribution of the fairly constant high - . 77. Meisenheimer's divisions based primarily on the limits of the tropical pteropods coincide in the North Atlantic with the 15 200 isotherm and in the South Atlantic 20 220 240 260 280 300 320 340 40 60 80 100 120 160 W. the boundary lines of these two to Meisenheimer the investigators differ essentially in that according and Indian oceans are of the Atlantic communities tropical animal to Ortmann. Ortmann 24 31 sets somewhat different (Fig.LOO. with that of 17 boundaries on temperatures in which the warmth-limited.. 120 100 region: heavily dotted line according to Ortaccording to Meisenheimer. whereas. Limits of the warm water mann. heavy broken line water. As shown in Fig. His boundary lines therefore bend far toward the equator along the west coast of South America and of Africa on account of the cold Antarctic currents and the upwelling of water from the cold ocean depths. they are united around South Africa.. entirely independent of the conventional tropics of and Cancer. lightly dotted line after Ortmann. heavy coast line. 77. in fact.

in the best-studied region. the North Atlantic. yet in the open ocean and apart from regions where opposed currents meet. other subdivisions can be made.328 Geographic Divisions of the Sea Both the warm and the cold belts of the sea can be subdivided. the smaller plankton organisms. and perhaps the higher temperatures are less rigidly selective than the low temperatures obtaining in polar seas. both distinguished by characteristic dominant animals. This allows the development of an extensive warmth-limcommunity. The speeding up of life ited stenothermal histories favors the appearance of mutations. relatively narrow strips can be located in which water temperatures and their accompanying animal communities change more abruptly than in adjoining regions. Such phenomena are particularly noticeable in terminating streams. but they apply also to circular currents. Pelagic animals are able to follow these shifts to some extent. the animal communities of the cold waters can be separated into arctic and boreal communities. Indeed. for example. but littoral animals are less motile as a rule and often exhibit special adaptations. This condition is especially noticeable among the pelagic animal com- . such as the well-known Gulf warm Stream. The location of all these boundaries shifts with the season. At any rate. and genmetabolism temperatures This is best seen in erations follow each other in quick succession. The warm waters of the equatorial belt lack that distinct seasonal periodicity found toward the poles. The boundaries between different temperature areas are usually not sharply defined either physically or biotically. Tropical marine communities The influence of temperature on marine animals has already been discussed and needs only a brief summary. Marine animals react to an equatorial belt of water with a temperature above 25 as distinct from cooler tropical waters lying on either side with temperatures from 20 to 25. Similarly there are recognizably different animal communities in the cold-water areas around the poles with temperatures below 10 as distinguished from those of the less cold waters with temperatures between 10 and 15. As a result of the continuous high and growth rates are accelerated. An equal mass of plankton at any given time would mean more food for the plankton feeders per week in the tropics than in colder water. the tropical waters are rich in genera and species but have a smaller number of individuals per species than are found elsewhere. streams augment their area in summer and are pushed back by the advancing cold in winter. as has been stated previously.

Tropical Marine Communities munities. which are entirely absent in cold water. separated by the continents of Africa and America. Among tunicates the Pyrosomatidae be- long to the warm waters entirely. the pelagic animal . the number of sea birds that secure their food from the uppermost surface water is so much less in the tropics than in polar seas. with few exceptions. are limited to warm water. When tropical pelagic communities are examined. The absence of flying fishes in cold seas well be associated with the adverse effect of too rapid cooling produced by the swift movement of the moist animals through cold air. the species of the cope- pod genus Copilia. The tropical plankton includes numerous free-swimming larvae of forms of echinoderms. Of mollusks. while only one or two dwarf species are found in polar waters. Thirty species of Appendicularia in the Atlantic. This may well be a basic reason why. 15 We have also seen that a different stratification of animal life occurs in the tropical as compared with colder waters. which accumulate toward the equator. most of the Sergestes. annelids. The absence of seals in the tropics may have been originally determined by this same phenomenon. 329 The tropical waters are therefore rich in index forms. Of crustaceans. as well as the shrimp Lucifer. as against 3 species (of 2 genera) in the cold. even among the invertebrates. occur in the warm zone. as do the salpas with few exceptions. Hydromedusae and the Charybdeidae live The Geryonidae among the among the Scyphomedusae only in warm seas. are present only in the warm water. while the communities of the colder waters may be distinguished rather by their negative characters. except in a favorable region such as that established by the cold Humboldt Current. all heteropods. The Siphonophora. and others. are confined to the tropics. the Atlantic and Indo-Pacific. and the upwelling water mass along the coast of Chile and Peru. and the latter are specifically different. with a consequent lack of concentration in the surface waters. It is worth noting that in spite of the complete separation of the may warm-water belt into two subdivisions. mollusks. representing 7 to 8 genera. most of the Euphausiacea. It is characteristic of the shifting boundaries of the warm-water belt that the northern limit for north in the northern flying fish extends about 5 to 8 latitude farther summer than in the northern winter. most pteropods. Finally. and the pelagic snails lanthina and Phylliroe. the flying fishes (Exococtus and its relatives) are strikingly characteristic of warm seas. it is found that about 20 species of Foraminifera occur there. particularly in that the depth penetrated by surface pelagic forms is greater.

Jan Mayen. more than a barrel of the scallop. For example. coris an increased size of egg (compare p. vitreus. 198). sedentary annelids in the Barents Sea. particularly in the Atlantic. * Corycaeus lautus. Michael Sars brought up more than a ton of the with a single haul of the dredge. specious. * Thus. Although the pelagic animals serve adequately to separate the tropical animal communities from those of cooler waters. the warm seas are particularly distinguished by the reef corals. The descendants places. part are identical closely related species that vicariate in the two situation similar exists among the schizopod and decapod a regions. often in larger a little to the southward. only bare rocks are to be seen. 7 papilligera. C. in the sea along east Spitzbcrgen. the southern limit coincides somewhat with Ortmann's southern border of the warm-water zone. f . and.! 2U most of the salpas. Geodia. C. Polar marine communities A direct result of the winter freezing appears in the absence of algae and of sessile animals in the littoral formations of the Arctic* Sea to a depth of 6 m. the boundaries lie within the limits of the tropical belt. The area of distribution of these corals is indicated in Fig. as is therefore remain in the immediate vicinity of their parents. plants and animals are present in numbers than in the adjacent boreal zone. and certainly the most abundant crustaceans. The significance of the connection of the two oceans in the region of Central America during Tertiary times in this regard is evident. but the northern limits are independent of it. especially among the benthic animals. Because of the low temperature.. are present in both oceans. They are limited in their distribution to a mean temperature of at least 20. 77. Below the limits of ice action. Ueterochaeta Like Corycaeus clausii and C. and their variety increases in higher mean temperatures. the (Scione lobata. at another near time. Pecten siliceous sponge. The suppression of free-swimming larval forms is a result. abundance. related with this fact on the other hand. thus enormous aggregations of well single species occur in many known in arctic seas. from which the drifting ice masses have scraped off all life.330 life Geographic Divisions of the Sea limited to warm water shows a great agreement in both areas. Calami* robustus. Thelepus cincinnatus) are found together in patches. and part of them are species. 'fecundity is reduced. for the copepod species occurring in both. vulgaris.

a fact at least Hensen summarizes the results of the investiga- by saying that the arctic areas are richer in plankton in the summer and the tropical areas are meager throughout the entire At any rate. Crangon antarcticus and Chortomus antarcticus. Antedon eschrichtii.* The polar plankton larvae. was similarly taken. in the Antarctic. The abundance of diatoms in the polar seas furnishes many plankton animals both in the north and the south with siliceous skeletal material. tions to date and only 700 in warm water above 20. long) in the Arctic Sea. latitudes. some of which appear in enormous numbers of individuals. The density of population of the pelagic animal life shows characteristic differences. other accounts catches of feather stars. still many genera and species find their tions for existence in these colder waters. Hydro- medusae are much more abundant in cold seas than in warm water. is swimming It also differs distinguished by the great scarcity of freefrom the warm-water plankton in its distinctive copepods. Though not so numerous as the warmth-limited types characteristic of the small. the great abundance of which is responsible in a large measure for the wealth of animal life in the Arctic. At the Gauss Station in the Antarctic. the volume * In the Arctic. . used. and great quantities of various species of pteropod snails among the mollusks. life At in least in the Atlantic. cold waters are richer in in general ~l Lohmann particular than warm water. such as the large Calanus hyperboreus (9 mm. obtained an average count of 2500 organisms in 1 liter and animal life in of cold water of value as a ratio. The number of animal species that can live in the surface water of both the polar and the tropical seas is The fauna of the polar seas shows throughout the influence of environmental selection as compared with that of the tropics. by many radiolarians (species of Aulocleptes) and 20 a number of ciliates (Latnprotintinnus). of the polar areas. tropics. groups are the Hydrothe coelenterates and the Holothuria among the echino- Siliceous sponges occur especially in antarctic seas. of the decapods.Polar Marine Communities groenlandicus. Sclerocrangon ferox. a very definite periodicity exists in the plankton year. On the other hand. Bythocaris and Hijmenodom glacialis. only a few species of shrimps are present in the higher 26 Hermit crabs and true crabs seem to be entirely absent. Scarcely less characteristic are the hyperiids and schizopods. the scarcity of higher crustaceans is striking. 331 mention large The composition of the cold-water fauna is different from that of the tropics in many respects. Such most favorable condi- zoa among derms. for example.

Hensen it is. Some of the mammals of the polar seas. on the other hand. especially the diving birds like the eider ducks (Somateria). In the North Sea. plant life is entirely wanting because of lack of light. a thorough light penetration of at least the surface layers for about 20 hours in summer must be extremely favorable for the development of the phytoplankton. and the former .000 in April. decreased in April to twenty times its lowest volume. while animals dependent upon plants survive only by reason of stored reserves. Whether the entire year's production of the polar sea is greater than that of the tropical cannot yet be stated with certainty. Thus. and thereby much food is provided for a rich fauna. and returned to minimum volume again in May. and only dormant stages survive. determined in large part by the strength and duration of the light. . like the walrus.000. The birds. these are lacking in the tropical seas or occur sparingly.332 Geographic Divisions of the Sea of plankton varied in the following proportions during the course of the year: it showed its minimum in winter (June to December). depend on benthic life for food. is inclined to believe that The abundance of the lower forms of animal life in the attracts upper a large number of bird and mammal predators. 700 walruses were killed in a few hours on Bear Island in 1606. Still others are active predators. and live mainly on cuttlefishes and fishes. feed on benthic animals. and the Weddel seal. increased to twenty-five fold in Feband reached its high point in March when it was fifty times greater than in winter. Better testimony could scarcely be given for the abundance of life in the polar seas during the summers than the immense numbers of these birds and mammals that found their sustenance there before their decimation by man. in- ruary. on islands and cliffs. and fully 900 the following year. but his evidence is inconclusive. Many. or in limited regions with special conditions. a vertical pull of a net through which 1000 tons of water was seined brought up about 400 Calanus in February and about 17 The enormous increase in summer is naturally 4. Foodstuffs necessary to the plants accumulate. in winter. creased sevenfold in January. during the warmer seasons. such as the whalebone whales or the crab-eating seals (Lobodon carcinophaga) which feed upon Euphausidae. others such as penguins feed on pelagic fishes and on plankton. according to Heuglin. Sharks occupy a strata of cold seas similar place in the food chain in the warm-water pelagial. others. live on plankton. are entirely dependent upon the sea for their nourishment. which breed in enormous numbers along the coasts of the polar seas.

each characterized by dominant forms. as is to cool and temperate Atlantic. the animal communities of while areas. Comparisons of the animal life of tropical and polar seas demonstrate marked be expected. 9 amounting in the southwestern portion of the North Sea to even on the bottom. and a temperate a boreal area with surface temperature of 2-8 area with a surface temperature above 8 have been distinguished. fairly closely allied to those of higher are not differentiated from those of tropical and latitudes. The relict hypothesis of Theel (1886) elaborated by . Comparisons of the animal communities of tropical seas with those of polar regions have been made primarily on the basis of the situation in the Atlantic. mean any northern and southern distribution disconwere focussed especially on the two widely separated populations. and existence of identical species in led to attempts to account theoretically for the present discontinuity 3G of their ranges. which are well discussed by Ekman. Such distributions led to an active discussion of the bipolarity tinuous across the tropics. and in the shallow coastal waters of less than 20-m. favor the development of eurythermal animals that may even invade the colder seas and thrive better there on account of the greater uniformity in conditions. Early lists of bipolar animals came to phenomenon of "bipolarity. with a temperature below 2. The greater temperature variations in the North differences. It would require too much space to consider these subdivisions.Bipolarity 333 vast abundance of the fur seals on the Arctic islands is one of the most notable of animal phenomena. where relationships are best known. In addition to the Arctic area in the North Atlantic. 5 13 In comparison with the equatorial Atlantic. the Indian and Arctic oceans and also the Antarctic have a greater development of coast line and are relatively - more shallow. depth to 15 or more. Even here the difference in density of population in warm and cold waters need not be related to temperature alone. 9 Bipolarity The investigations of marine life stimulated by the Challenger Expedition (1872-1876) disclosed what seemed to be a surprising number of animal species found in Arctic and Antarctic waters and unknown from intervening seas. sharply subtropical waters. but. Unusually large amounts of plankton from the tropical Indian Ocean are reported." By extension of ideas. Similar relations prevail in the East Indian Archipelago and undoubtedly affect the development of pelagic life.

on the reported 1898 40 ) facts of species bipolarity. which are reported also from New Zealand seas. the Cretaceous seas." phenomenon series of Turning to bipolar genera. the relicts in the sense of Pfeffer and the results of "migration" in the sense of Ortmann. Myriothela. opposed the structure of hypothesis and D'Arcy Thomp- son ( identity. and Sclaginopsis. Grammaria. with subsequent evolution predominant with survival of the 30 Ortmann built (1896. and it was this continuous connection of the two polar seas through the colder water at considerable depths in the equatorial zone that had been pointed out by Ortmann in his socalled "Migration Hypothesis. . 327-329 ) of remains to be proved out that actual range disjunction points for even this list. Since the turn of the century. Even in groups that are widely distributed there may appear geminate species that are respectively Arctic and That the modern Antarctic. The unsatisfactory nature of the examples of bipolar be bipolar is critically examined by Ekman " . the euphausian Thysanoessa. there have been continuing discoveries of supposedly "bipolar" animals at greater depths (and absent at the surface) in equatorial waters. and thus ancient forms in the waters of high latitudes. Didemnum albidum and Botryllus schlosseri. the gephyrean Priapulus. These have reduced the necessity for a relict hypothesis. by the North Atlantic ascidians. Such examples recall the isolated occurrence of the horseshoe crabs in the western North Atlantic and forms cited at the species level is illustrated the tropical East Indies rather than any distributional properly to be referred to as "bipolarity. the sardine Sardina. pp. it is quite evident that a genuinely bipolar distribution is demonstrable in a conThe examples cited by Ekman are the siderable series of forms." The residual series of species thought to who ( 1935. and the entire group of stalked medusae. trachymedusae Ptychogastria and Botrynema. furthermore. Antarctic species form what in is which a series of Arctic and a evidently a natural group. It presented a sharp critique of the suppositions of specific could be shown that in most of the forms in question the evidence of identity was extremely inadequate. Hydroid. as in the hydromedusan Bougainvillia. distributions an earlier separated represent continuity is evident from the forms are thus both and resultant any evolutionary consideration. the order Stauromedusae.334 Pfeffer 33 Geographic Divisions of the Sea (1891) explained the known discontinuities by the supposition of world-wide uniformity of climatic conditions and of animal life in in the equatorial or low-latitude seas. the cephalopod Bathypolypus. 1899 32 ). genera Lampra.

bays.3 in the middle part of the Red Sea and 35. Temperature distribution in small seas frequently differs greatly from that of the ocean. although the highest surface temperature always occurs in the lesser seas. Animal communities of As stated at the seas. The peculiarities of the lesser seas rest primarily on their salt content. and gulfs beginning of the present chapter. crease in salt content as a result of increased evaporation. 203). but from other. proper. Bays. bays. the decrease in temperature according to depth is 18 . in the polar seas. the phenomenon is certainly one of "bipolarity" and one of essential interest to ecological animal distribution.Animal Communities of The phenomena Seas. 79.6 in the Persian Gulf. Finally. may equally as well be indederived from the pendently intervening warm-water form or relict of the form from which the warm-water form populations primary parallel cold-water "varieties" that has evolved. The salt content of the ocean and from ice and of water melted of various bays and land-locked seas has been given previously (p. similarly characterized subdivisions. in the long The parallel development of great masses of plankton summer in Arctic and Antarctic seas. salinity. If the deep-water equatorial forms are found at the sur- face in both the Arctic and Antarctic. and near coasts and in the western Pacific 32. not only from those of the ocean ocean. which is evaporation. the physical conditions of the partially separated seas. in to whereas in the open ocean the summer temperature some places rises to 29. and is summarized for the Baltic Sea by the lines of equal mer because of the small salinity in Fig. with complex atis not less interesting for the fact that the seals of the two regions belong to different genera or that there may be distinct types of whales. and Gulfs 335 of bipolarity have an ecological aspect somewhat of the systematic relations of the species or their geoindependent logical history. reaching 34. and gulfs are associated with the lessened interchange of their water with those of the main Important differences in temperature. on the other hand. there remains the development of tendant food chains. The temperature of the surface layer is not much different from that in the neighboring part of the ocean. and other conditions are found in each such sea or bay. Further. such differences tend to separate its animal communities. a decrease in salinity takes place in the sum- regulated by the ratio between influx of fresh water and The tropical areas of the ocean show a noticeable in- amount of evaporation and the huge amount snow.

Furthermore. tide__ movements are often very small. The two bodies of water are quite unlike hyrographically in that the Gulf of California is in connection with the adjacent ocean down to nearly 3000 m. e.g. high temperature controlled by the depth of the containing slight 38 The movement of the deep waters of many seas caused by is the shallowness of their connection with the ocean leads to the further result that the oxygen content accumulate.. But if the entrance to a deep sea such as the Mediterranean (see Fig. 15) is closed for the deep water by a bank or shallow strait. though it opens farther to the north. in the Sulu Sea. In water from the ocean can enter Gulf the consequence deep tropical of California. 30 and the small number of mussels and snails 3T in the depths of the Red Sea is striking. In the depths of the Black Sea. the life animal Animal life is Thus the lack of ptychobranchiate ascidians. may in the deep parts of the Mediterranean. The Gulf of California is somewhat comparable to the Red Sea in general climate. the flood tide for the Red Sea is 1-2 . the cold water from the depths of the ocean cannot enter and the temperature of the restricted sea bottom remains constantly at that of the lower layer of the inflowing ocean water. so that from that depth to the bottom all life is absent. to its greatest depth at 2190 m.336 different in Geographic Divisions of the Sea many small seas from that in the ocean. low and that substances lethal to In the deep basins of the Baltic Sea. therefore sparse. The low tempera- water masses is explained by the slow creep of the ture of the abyssal^ denser polar waters along the bottom.. Similarly. the temperature of 21. especially of the Levantine and Ionian seas. whereas in the ocean at such depths the average temperature is 2. Basins within the Gulf follow the rule documented in the preceding paragraph of having bottom water with the relatively sill. in the seas. the unusual scarcity of Hexactinellida (only two species). to the bottom (more than 4000 m. and support a rich animal life. whereas the greatest depth of the sill at the opening of the Red Sea is only about 100 in. 2r> gen sulphide. This is not universally true. per liter even at 180-m depth. there is a considerable amount of hydrois water poor in oxygen and rich in carbon dioxide. with the exception of anaerobic bacteria. In the Red Sea.3.) remains at 10. shape.33 cc. and distance from the equator. and apparently also in the depths of the Red Sea.11. whose connecting straits are only 50 m. amounting to 0. the temperature from a depth of 800 m. come to the surface towards the north.5 extends from a depth of 700 m. deep in the shallowest part..

1 - In the Caribbean Sea a characteristic deep-water fauna has littoral developed from descending species.- warm. the formation of species is favored by the isolation of seas in conjunction with the peculiar conditions existing within them.. disappear comof starfish that go down into the pletely.g. Finally. on accordingly very small in these seas.6 cm. in the Baltic at if 337 amounts to scarcely the average. It is therefore to be expected that these seas should have species.The Mediterranean Sea m.. In the Red Sea the number of endemic animal species known to 1918 was: 1 hexactinellid. The temperature barriers. to8 ptychobranchiate ascidians. such as the Mediterranean and the Baltic. 16 further investigation of with gether ranges in near-by waters may decrease these numbers. 17 endemic species of macruroid fishes. whereby stenothermal. corresponding with representative species in the neighboring in spite of its and often very seas. which as surface layers of the ocean. 4 crustaceans.. although otherwise there is marked similarity in the two populations. on the other hand.water animals greater than in the stay in the . In the Mediterranean. as they find much less food along such a coast than along a shore whose tide flat is more extensive. The Mediterranean Sea Only a few seas have been adequately investigated as yet. which become more numerous with The Mediterranean and Baltic seas will be discussed below in this connection. and this has its effect upon the distribution of the intertidal inhabitants. and at Jasmund is it is 3.5 m. are among the best-studied areas. both in similar depths of salinity and bottom temperatures are higher than the reasons these Atlantic. These may therefore be further considered at this point examples of the peculiar animal communities in their special environment. in the Mediterranean. shore birds are much scarcer. For the neighboring abyssal animal com- marine as munities also differ from those of the connecting ocean. but some. primarily because of the shallowness of the Straits of Gibraltar and the low ratio of inflow to evaporation. 35 bivalves varieties Memel 1 cm. 21 snails. The Sulu Sea (24 littoral small size contains a significant number of endemic distinct species. and 11 from the deeper parts). Thus the number of species lightless depths of the Mediterranean 2 is much Alciopidae. live at great depths in the MediterAtlantic. e. for example. warmth-limited animals are prevented from entering greater depths. 0. The tide flat endemic and increasing isolation.

This applies. have been obtained down to 1200 m. Mediterranean depths. an active descent of the 8 surface water into the deep has been demonstrated. But the deep water of the Mediterranean does not everywhere contain sufficient oxygen to permit active animal life. Scrramis gigas. salpas. and many others. a number of fishes com- mon in summer Julis turcica. crease is The an unsolved problem why many animals in numbers present pelagic large directly west of the Straits of Gilbraltar do not occur in the Mediterranean. 11 Straits of Gibraltar life Conditions in the western part of the Mediterranean nearer the and the open ocean are more favorable for animal than those in the eastern half.338 rancan. limited in the Atlantic to the lighted layer.i0 Geographic Divisions of the Sea heteropod and pteropod snails. is still example. for unknown.. sink to the bottom. can survive in the Mediterranean. at Naples. and die with further decrease in 19 Such forms can survive the winter in the warm temperature. to the fishes Argyropelecus olfersi and Vincignerria sanzoi and to a number of eel larvae. . Experiments in the Naples aquarium show that these animals do not tolerate a decrease in temperature below 12. Appendicnlarians. which do not ordinarily descend deeper than 300-350 m. which are carried toward the Straits of Gibraltar by the southern arm of the Gulf Stream in some numbers. many heteropods. the ocean sun- Mola mo/a. also at times found in masses (Salpa maxima) surface are animals. are found below 1000 m. chiefly bottom nets. and in the Aegean. Such forms include the scyphozoan Charybdea. many siphonophores. salpas.. Though in some fish places.. the turtle Caretta caretta. starfishes. they become inactive.. e. whereas the larvae of the river eel . In the Gulf of Naples.g. in the northern part of the Balearic Sea. also descend to a depth of over 1000 m. Pyrosomas. Thus tropical and subtropical forms. and prosobranch just as it and many animal groups such as snails have fewer species in the exact cause of this eastern de- eastern than in the western half. in other parts therefore so poor in animal life that Forbes* early investigations of this area led to the erroneous conclusion that the depths of the oceans in general were is the deep water rich in carbon dioxide and is without animal life. lacking in the ocean below 400 m. in the southern part of the Adriatic. the tropical jellyfish of the genus Lirtope. e.g. are not caught in the cold season. 1 the contents of fishermen's the among - The high temperature of the deep water makes it possible for many stenothermal animal species of warm water to survive the winter in the Mediterranean.

so that it may be assumed that a large number of those species do not occur in the neighboring seas. in length compared with 26 cm. and the inflow is at deeper levels. are fairly well known. and 4 species and varieties of Ap- Of fishes. great influx of water is relatively many respects. benthic and known only from the Mediterranean. the barnacle Pachylasma giganteum. perch. may be considered endemic only with reservations. Clupea papalina is the mediterranean form of the northern sprat (C. As the outflowing water is less dense. the anchovies (Engraulis encrasicholus) pendicularia. whereas the inflow from the ocean is much less. and from the Baltic relatively large quantities of water flow outward into the ocean. among others. 21 decapod crustaceans. contain a race distinctive of the Mediterranean. on account of the lower salt content.The Baltic Sea 339 "Leptocephalus" brevirostris pass through the Straits of Gibraltar in 35 large numbers. The salt content. yet in The In Baltic Sea is just the opposite of the area land sends its waters into the widespread small and shallow and the basin. and the sardine (C. jonstonii). are Species of the most widely separated animal groups. 2 in the Baltic Sea basin. and several other fresh-water fishes have been caught in the Strela Sound between the Island of Ruga and the mainland. is greatest in the western Baltic near the connection with the ocean and distant from the great inflow of fresh water. In this category belong. as well as of the Azores and the Canary Islands. sprattus). even of the surface water. and a few from fresh water live in the Baltic Sea. 78). the outflow occurs on the surface. The number of species of marine animals decreases regularly from the west toward the east. As a result only very euryhaline forms. 9 in Kiel Bay. brackish-water animals. the Baltic Sea 3 * 4 ' 23 . pilchardus) never attains the length of the oceanic form of this species These species the faunae of the French and Spanish coasts of the Atlantic. For these reasons the salt content is small. A accompanied by the smaller loss through evaporation in these latitudes.34 Mediterranean. and only 1 in . and decreases toward the east until a salt content of only 2 c ff and less is reached in the / surface water in the Gulf of Bothnia (Fig. the opposite of the Mediterranean situation. (maximum of 18 cm. two species of starfish (Astropecten spinulosus and A. pelagic. and the number of fresh-water species rises in the gulfs of Finland and of Bothnia.). 2 species of salpas. thus 55 species of decapod crustaceans are found in the Kattegat. Pike.

7% seems to represent a limit that even euryof . the western Baltic 9. Copepod and individuals become fewer toward the east. 14 Differences also occur in the plankton. and to these may be added Bosmina maritima and farther to the northeast a large number of rotifers. After Brandt. and only 2 species extend much beyond the east coast FIG.340 Geographic Divisions of the Sea the Gulf of Finland. while the cladospecies cerans Podon and Evadne increase. the The salt content per thousand in smaller vertical depth in meters in larger oblique figures. 78. the Baltic basin 5. of the annelid genera. Map of the Baltic Sea. Sweden. figures. Nephthys and Nereis. The salt content of 3. the North Sea contains at least 20 species.

is present at . from Skagen to Lulea. 6 of crustaceans. Perhaps for the same reasons that bring about their decrease in numbers FIG. But in a few limited places (see Figs. 36 cc. 6 species of worms. only the barnacle Balanus improvisus and the bryozoan Membranipora pilosa extend beyond it. 4 of bivalves and 1 of bryozoans. while only 32 cc. depth 2 species of worms. occur. and at greater depths than 230 m. of species. the Baltic animals decrease in size the farther east they live. and at 200 m. according to Ekman. After Brandt.. of carbon dioxide per liter is present at a depth of 0-60 m. only one-fifth. the copepod Paracalanus parvus. 9 are here unfertile.. in Longitudinal section of the Baltic Sea. 79. Harmothoe sarsi and Priapulus caudatus. only the sprat (Clupea sprattus) and the needlefish Nerophis ophiodon propagate in the Baltic. has a depth of 427 m. The reason for this condition lies in the low oxygen content and the accumulation of carbon dioxide. depth toward the east. depth is about one-third saturated with oxygen. the average depth amounts only to about 67 m.The Baltic Sea 341 haline sea animals pass with difficulty. On the other hand. depths in meters at the left. located south of Stockholm. the most important of these depressions. but in spite of this. and indeed only those that occur also in shallow water. only one-fifteenth. e. which increases in At depths of 80 m. only 17 species of bottom forms are found. isohalines. in the deeps of East Gotland at a depth of 100 m. 28 Of 11 species of marine fishes. they contain only an impoverished fauna represented by few individuals The Baltic Sea is in general a shallow sea. which are common to the Gulf of Finland and the North Sea. 78 and 79) the bottom drops off to rather great depths. all animal life is absent. more than 150-m. of small size. with % c . In these depressions. Many immigrants from the North Sea are not able to propagate in the Baltic. and they restore their numbers only by means of ever-renewed reinforcements. the salt content seldom goes below 12%<?.g. The water at at 0-60 m.

Komm. Fischer: vi. It has retained a number of relicts from its connection with the White Sea by means of Ladoga and Onega seas. peculiar development of the animal life in such seas shows how obviously the composition of animal communities depends upon en- The vironmental conditions.. on the occasion of continuous easterly storms. Aus den Tiefen des Weltmeers. 390 text figs. "Die zoologischen Arbeiten der Kieler Kommission 1870 bis 1920. the Skagerrak. insbesondere die der Kieler Bucht. 1887. "Die Fauna der Ostsee. 594. 7 pis. e. which is represented by water lakes. It is because of this geological youth of the Baltic that the num- ber of endemic species is small. Preuss. the Baltic is still young. Halicryptus spinulosus. 2 maps. Karl. Idem. and the Kattegat. Schilderungen von der Deutschen Tiefsee-Expedition. C. and 41 Geographic Divisions of the Sea cc. Plankton Exped.. 76-194. 5 pis. 4. Idem. B. at 200 m. and the study of the effects of the freshening of the waters of the eastern Baltic Sea makes a suitable transition between the consideration of marine animal communities and those of the fresh waters.g. BIBLIOGRAPHY 1. the surface water recedes in the east and an undercurrent of water rich in oxygen and salt advances farther toward the After such storms east. 1900. 14 text figs. 1921. Heft EaB: 1-68.. 1897. is found in the northern seas but is absent in the North the M . 2.. Sea. "Die Thaliacea der Plankton-expedition.. 1 pi.. 27 were taken in the Danzig deeps in February." Bibliographic Zool. which are treated in the following section. Idem. thus. but is lacking 10 so also Mi/sis in the North Sea.." Festschr. 2 maps (p. Carl. "Die Alciopiden und Tomopteriden der Plankton-Expedition/' Ibid. richer animal life becomes possible in these local deeps. species of annelid normally occurs in the western only Geologically speaking. Apart from a few races of fishes. 2. Chun.. 7 maps. relicta in the neighboring freshoculata. Jena. 7:10-34. . 1:1-66. Meeresunters. Heft Hb: 1-61.. bringing fresh supplies to the depressions. 1 while 1904." Verhandl deut. 3 species of maldanids (Annelida) Baltic. 5." Ergeb.342 100 m. Zool Ges. which likewise occurs at Spitzbergen and the White Sea. Brandt. 3.. 368). pp. Rarely. Salpen. (p. Wiss. "Die pelagische Thierwelt in grosseren Meerestiefen und ihre Beziehung zu der Oberflachenfauna... 24). the starworm. 1900. Vertheilung der 2. 1894. 6. the copepods Pseudocalanus elongatus and Temora longicornis may be mentioned as endemic. Apstein.. 46 pis.

542. Hubbs.). 1:259-351.. Ergeb. Lohmann. Kiel. "Die Bevolkerung des Ozeans mit Plankton. 3. JR. 153.." 9.f.und Ostsee unter Beriicksichtigung von Formen des Nordatlantischen Gebietes. Johannes. Gilbert. 1935. 20. "Einige Abt. einschliesslich der verbindenden Meeresteile. "Notizie biologiche riguardante specialmente il Mitt.). Murray. xii. 27-31). Forbes. "Die Pteropoden der Deutschen SiidpolarExpedition 1901-3.und Pflanzenlebens in den von der Deutschland wiihrend ihrer Fahrt nach Buenos Ayres durchfahrenen Gebieten. "The macrouroid fishes of the Philippine Islands and the East Indies. Hubert.und Ostsee. "Critische Uebersicht der Fisch-Fauna Finlands. a general account of the modern science of oceanography. Kiel. 22. 3 pis." New Philos. Richard." Bull. Biontologie.). H. 18. based largely on the re- . 1906. Abt. 9 (Zool. vol. 24. F.. map (p. Meeres. (n. A. Planktonstudien. Jena.. being the substance of a communication made to the Royal Institution of Great Britain. Monaco: 123-129. 9th Intern. John. (p. Ekman. d." Wiss. (n. pt. 1892. 1. Adolf. 204 ff. U. 1902. Sta. Handworterbuch Naturw.f. 100. and C.. 3). 1864. 3:1-617.." Wiss. 19:513-761. 17 figs. 1895. C. 1). 5 maps (p. 121). 1914." Arch. (p. Heft 2:9223. Meisenheimer. Naturgeschichte. 135 17." fig.. "Die Sipunculoiden der Nord. Edinburgh. 20 text figs." 8. 343 Copepoden im Ocean.. figs. vol. (p. 2:461-744. Jena. 21. 30. 26.. Haeckel. Hesse. and Johann Hjort. 1894. 105 (p. C. 14. Fischer: viii. figs. C.. Lo Bianco. 1912. L. Meeresunters. Rev." Salvatore. Abt. "Les expeditions oceanographiques danoises dans la Mediterranee. 244 figs. Hydrobiol. 6:789[Article] "Meer. light Edward. 723)." Bronns Klassen und Ordnungen. Sven. 15. 113 text figs. Friedrich. Akad. Krummel. Zool. Museum.. Idem. (p.. Wiss. Drechsel. 1 1924. 801)." 1 W. Arch. Zool Ges. H onsen. 68 ff. Fiskeri og Hvalfangst i det nordlige Norge (pp. 1 pi." 1909. J. 113ff. 12. 1:369588. des Atlantischen Ozeans. 613.. J. Johannes. 11 pis.Bibliography 7. 1912. 1 pi. Malmgren. 40 figs. 11. the 23rd of February. Zool Congr.).. Natl. 19.." 4. Fischer: V. Ludwig. 72 ff. Ergebnisse der Expedition. "Ascidia Ptychobranchia und Dictybranchia des Roten Denkschr. 4:61-80 (p. Johann." Int. Otto. Tiergeographie auf oekologischer Grundlage. "Ueber die horizontale und verticale Verbreitung der Verhandl.) 16:85-127. 1920. 1914. 74). and Otto Hamann. Fischer. 1920. Hans. 1918. S. Exped. Hjort. 1844. 1890. Friday evening. 25. 1911. Leipzig. "Die Nephthydeen und Lycorideen der Nord.). "Beitrage zur Charakterisicrung des Tier. Plankton 16. 95:1-123. 16 pis. Siidpolar Exped. Meeresunters. 1844. "Echinodermen (Stachelhauter). 10. 815 (p. 36:318-327." Ergeb. 5:185-225. Akademische Verlagsgesellschaft: xii. 1912.. 14).) 13:1-87. 2. Tiergeographie des Meeres. Heinen. A: 18-46. Ernst. Wien (m. The depths of the ocean. d. Neapel. "On the 13. Michaelsen. Dahl.-n. (p. thrown on geology by submarine researches. periodo di maturita sessuale degli animali del golfo di Napoli.

Fisher: iv. H.) 15:1-94. With contributions from A.und Ostsee. physics. Prentice-Hall: x. 35. 22:311-349. 9:571-595... 1-62. W. Kiel. 1914. Theel. Helland-Hansen. 1 text fig. 7 charts. F. A. (n.. Pfeffer. 1:1-36.. "Die Metamorphose der Plankton-Copepoden der Kielcr Ibid.. Idem. 1893. 27. 3 pis." Nature. M. 1896. mit besondcrcr Beriicksichtigung der Dekapodenkrebse. Ibid. 2. U.. 5 pis.344 Geographic Divisions of the Sea searches of the Norwegian steamer Michael Sars in the North Atlantic.: 120 pp. Nolte. jetzigen Verbreitungsverhdltnisse unserer Tierwelt." Zool Jahrb. 40. Proc." burgh. 7 pis. figs. "On a supposed resemblance between the 39. 8 text figs. Appelof." Ergeb. Sverdrup. Wien H... Thompson.. 33. 30. Y. their N. "Dekapoden und Schizopoden. Heft Gb.. 32. The oceans. 1899. Edin- . Ortmann. 96. Schulze. R. 70). d. E.. Johannes. Gran. Sturany." 69:255-297. Ges. and sipunculids dredged by the Swedish Antarctic Expedition 1901-1903 and the phenomenon ofr bipolarity. Versuch iiber die erdgeschichtliche Entwicklung der 31. Hjalmar. Vetenskapsakad. Jena.-n. Syst.. Oberg.f. London. 1896. Verhandl. "Priapulids marine faunas of the Arctic and Antarctic regions. chemistry. Soc. 47. Idem. 265 biology... D'Arcy W." Am Naturalist. 33:583-591. H. Idem. Hoy. (p." Rothen Meeres. Die Bodenfauna von Nord- Hamburg: und Ostsee. 93). 1 map. Bucht" 29. "Uber Bipolar! tat in der Verbreitung mariner Tiere. 1900. illus. Macmillan: xx. Georg. "Hexactinelliden des Denkschr. J. Schmidt. HandL. 38. 109:45-47. 1922. and B. Plankton Exped. 9:37-103. E.." Kgl.. Fleming. Svenska. 1911. W. A. 24:221-235. 69:311-324. "On new facts lately presented in opposition to the hypothesis of bipolarity of marine faunae. H. Max. Maldoniden der Nord. Akad. 37.). and general Johnson. 3 maps. 821. Wiss. No." Wiss. Meeresunters. Reibisch. and R. 1901. 1942. 1891. Grundzuge der marinen Tiergeographie Anleitung zur Untersuchung der geographischen Verbreitung mariner Tiere.. 34. (m.. 7 pis. "Oceanography of the Gibraltar region. 36. 1898. Wilhelm. "Zur Kenntnis der 28. "Lamellibranchiaten des Rothen Meeres. 1087. 7 pis. 1906. Zool. 1 map (p.

Inland A Phase of Limnology .Part 3 The Distribution in of Animals Waters.

.

rivers. of which only a few species are found. especially the difference in number of individuals and species. The ocean covers immense portions of the earth's surface and is one continuous body with only partially separated arms or seas. The Environmental Factors in Inland Waters THE FAUNA OF INLAND WATERS DIFFERS IN MANY IMPORTANT RESPECTS from that of the ocean. size. and inland seas form innumerable bodies of water of spread over the continents as islands are in the sea. Inland waters. the Nemertinea. Whole groups of animals. e. and the Bryozoa. and many others have fewer species. The greatest differences between the environments represented by oceanic and inland waters relate to space and time. Gastrotricha. gephyreans. and tunicates. in rapid succession. or are essentially restricted to lakes and rivers. ciliates. comparatively as a rule are of comparatively short duration and appear and disap- pear. such as echinoderms. As has been stated before (p. as are amphibians. or at most separated into a few large divisions for short periods of time.. geologically speaking. which are often less rich in individuals: for instance. the annelids. is that the living conditions in inland waters vary so much from the optimum that they demand certain definite adaptive adjustments that relatively few gill-breathing types have been able to acquire. 347 . which are plentiful in the ocean. whereas the fresh-water lakes. the Coelenterata. and rotifers. Only a few groups are more plentiful both in number of individuals and of species in inland waters than in the ocean. on the other hand. brachiopods.g. The ocean has probably more system been in existence as one body of water since the beginning of geological evolution. are wholly absent in inland waters. the reason for these differences. each every or less isolated from the rest.It). squids. 43).

Only 17 lakes and seas have a surface area more than 10. even the deepest are shallow. compared with the ocean. In streams whose contents change rapidly and whose tributary area. they contain relatively little dissolved material. km. Only a few are deeper than 400 m. fresh when they^ into flow other streams. so that its variation is always within narrow limits. have chemical constitutions of their own.360 sq. contains only 0. The variation of the dissolved substances of inland waters is great. is 0.8% . The water of the Rhine. the extent of shore line and bottom in relation to amount of water is very much greater in inland waters than is it is in the ocean. km. but the ratio is often much it is seen in the Aral Sea where 10. on the other hand. the majority reach a depth of less than 100 m.. 1706 m. even in their deepest part. but where standing waters are fed the salt amount soon become Nevertheless. such differences are equalized.). or in the salt . Inland waters are usually fresh waters.. which has an area of 438. Lakes. especially.000 sq. and Lake Tanganyika. km. in greatest depth.3%o. compared with the waters they contain. salt lakes are formed. the ratio of extent of suitable habitat for sessile forms and free- swimming forms Salt content very different. depending upon their substrata and their tributary areas. Most inland waters are much smaller. to be sure. With the exception of the Caspian Sea.) have a depth of more than 1000 m. while only two inland waters (Lake Baikal. and. Lake Superior is about 305 m. 1435 m. but are present even in such waters.14 part of NaCl to 1000 parts of water.. is less than in the ocean.000 sq. They are usually of no great extent or depth.. Nearly inland waters.000 sq. the differences in the amount of dissolved material are constantly being all equalized. is widespread.348 Environmental Factors in Inland Waters restriction of space The becomes evident in the extraordinary varia- tion in size of inland bodies of water. km. for instance. and so. i. ponds. and pools decrease in depth until they merge into marsh. The depth of inland waters also varies. as a result. and. and we find all gradations of size of ponds and puddles. (Lake Superior has 82. There content rises. no lake or inland sea has an area of more than 100. The content of common salt. as is ratio for a salt lake aided by evaporation. in consequence of the circulation of masses of water.e. The average depth of the ocean is about 3795 m. The minimum higher. many inland waters contain a considerable are salt springs whose waters... In the ocean. a good of salt. The same to the down Consequently gradation of depth is noticeable in running waters..

and rather low mineralization. but not in the Jura or in the limestone Alps. Fresh-water sponges. 3 Victoria Nyanza has been ascribed to the want of lime in the water. Pisidium. the salt content changes with the seasons. Variability of lime con- and the composition of the fauna changes tent makes little very fresh waters very different from sea water. occurs in the lime-rich lakes of the Aland Islands but is not found in the inland waters of Finland u The smaller size of snails and mussels in that are poor in lime. Pisidium has very thin when growing under such extreme conditions.0 p. large. In the same lake region. for example. rich in lime. of dis- solved carbonates. Fresh waters also vary among themselves in the content of other salts. the finger-nail clam. A wealth of lime in water is favorable for the development of snails and mussels. correlated with the mineral content of the water. rains freshen the water Sea where In the salt seas of the steppes.Salt Content where it is 349 swamps of Laneuveville in Lorraine it is 54% . have shells so thin and poorly mineralized that when 16 fresh and wet they can be bent through almost 20 without breaking. growing in extremely soft and acid waters. high organic content. Bryozoa.1 to 6. 20 Holopedium is most commonly found in the mountain lakes of Europe. acid waters. in consequence. are intolerant of excess lime. although found under other conditions. or in the Dead 231. Iceland.m. for For that reason we find no instance. and Campeloma. well-developed specimens of Pisidium have been taken. the distribution of fresh-water sponges is. the gilled snail. are characteristic less dissolved with soft waters that have even shells forms in clear lakes carbonates than the typical bog-lakes with colored. 16 and in west Ireland the Spongilliclae or Bryozoa in Montenegro. Mussels of the genus Anodonta. and hard waters.p. attains its best development in small lakes of dark color. This lime content is a decisive environmental factor in animal distribution. high in waters flowing over dolomite or limestone. which contains calcium carbonate. especially of calcium and magnesium. where periodic and long periods of evaporation during the dry season concentrate it again.1 part per million.3%o. and America. poor in lime. with a pH of 5.0 to 5. We distinguish between soft waters. The calcium carbonate content is low in the waters of granitic or sandstone areas. mollusks are present even in acid water with a pH as low as 5.1 and with 3. mountain lakes are much richer in sponges than the limy lakes. In water in which the content of silicon . while in some bog-lakes. and the cladocferan Holopedium gibberum. Ncritina. to some extent. In the Highland Lake District of Wisconsin.1 and a calcium content as low as 0. Spongilla lacustris.

even sometimes to warm springs. this is the more interesting in that these spicules have been regarded as an important species character. and the waters in rivers flowing through extensive swampy virgin forests are especially rich in humus. The waters of the North America afford an example. which gives them a dark. bordering the site of the former lake. lacustris low in total solutes. S. Hydrogen sulphide as in landlocked Organic content Still by their content of organic matter. because of the H L>S content of their waters. shows a progressive at- its spicules. 7 theless in inland waters is just as deadly to animal life arms of the ocean. rivers as well as lakes. for instance. Simi- tenuation of larly Tubella pennsylvanica shows decided variations correlated with the degree of mineralization of the water.4 in Inland is Waters mg. and the regional salmon (Salmo coregonoides) is not found in the main stream but only in the mountain tributaries. is present in solution in the Puga creek in Ladakh (Kashmir). the microspinal dermal spicules are lost. Borax. The surface animals that descend into the depths periodically or whose winter eggs sink to the bottom are at a disadvantage. High humus content makes such waters uninhabitable for many animals. blackish brown color. per liter and which also as the mineral content decreases. snails. Crustaceans. and fishes were diminuIn regions of alkaline flats remaining from the evaporation or tive. especially when frozen over. drainage of formerly extensive fresh-water lakes in the American Southwest. Crustacea and fishes are never- found there. A contamination of inland waters. The Obi River as it flows through the plains takes up so much dirty steppe and morass water that it becomes stagnant and foul. 8 some cases these abrogate accepted generic Other inorganic compounds are also dissolved in great quantities in many inland waters. but in which the animal life was inhibited by this factor. often occurs near the settlements of man. If the water is sufficiently soft. Dismal Swamp in eastern . the fishes and other life may be restricted to springs. some Swiss lakes are unfit for animal life at depths below 13 m.350 oxide is Environmental Factors below 0. so that certain migrating salmon that are found east and west of the Obi do not enter this river. and the* multiplication of the name "Rio Negro" on the map of South America reflects this phenomenon. 6 Sarasin found a lake in New Caledonia whose waters were very rich in iron salts. other inland waters. are characterized Moor and bog waters and the lakes fed by streams from such sources. especially through organic matThe degree of conter. in criteria.

and consist species of animals thrive. the oxygen content is reduced and the hydrogen sulphide.. in practically pure water mayfly nymphs. and carbonic acid content increased. This is. Finally.. the mesosaprocoles live: very many protozoans. The presence of salmon and lake trout (SaZmo lacustris) is also regulated 6 by the oxygen content of the water. The oxygen content of inland waters is also subject The water of mountain streams is well aerated to much varia- in the spray of rapids and waterfalls. The deep water of many lakes is as rich in oxygen as the surface water. in lakes with a thermocline. In the lower reaches of rivers the amount of oxygen is less and depends upon the number of oxygen-producing plants as well as upon the amount of contamination from tributaries. The effects of pollution by city wastes in certain American rivers are discussed in the final chapter of this book. of course. and crayfishes are found. both in great numbers. etc. in the Alpine lakes and in the clear Eifel crater-lakes. however.1 In standing waters only the relatively small. mostly especially flagellates.Oxygen Tension 351 tamination can well be estimated by the composition of the fauna. stonefly nymphs. smooth surface is involved in the absorption of oxygen from the air.. per 24 liter. e. Oxygen tension tion. during the summer. numerous worms and rotifers. of protozoans. and some lower crustaceans appear. as well as a number of species of fishes. iron sulphide.8 cc. the mussel Sphaerium corneum. for they are found only in waters with a high oxygen content. the isopod Asellus. Thus Lake Nantua in the French Jura . in others. In the Elbe.g. near Hamburg. per liter. e. and oxygen is being given off by green plants. Farther down from the origin of contamination.g. and at 18 it may contain over 9 cc. Through the disintegration of highly complex organic compounds. and finally a number of insect larvae. the deep water is poor in oxygen content. On the other hand. the oxygen content fluctuates between 4 and 8. In such waters only a few These are called saprocoles. of the greatest importance for the animal life on the lake floor. where the albuminous substances have begun to be transformed into amino acids.5 cc. of Metazoa only the slimeworm Tubifex and the larvae of the drone fly Eristalis are generally present. Farther down in comparatively clean water some of the oligosaprocoles. among thick Potamogeton or which is the usual amount at normal saturation at 18. the water may be oversaturated with oxygen dur- ing the daytime because of the production of this gas by green plants. Gammarus.. especially in shallow lakes with muddy bottoms or Spirogyra instead of 6.

in contact with which somewhat found. is in itself a shelter that slows further cooling. but the temperature of the deep water remains at 4 ice covering. seldom freezes to the bottom.000 gram-calories per square centimeter of the lake's surface. When the surface waters cool they become heavier than the warmer waters beneath them. wide and with a mean depth of about 30 m. and sink while warmer waters rise to the surface. This determines a condition of greatest importance for the survival of fresh-water animals during the cold season in the temperate and frigid zones. Further cooling makes the water water does not sink as it cools below less dense. Lakes of eastern United States about 10 km. which are more marked. so that the surface 4. Temperature The temperature of inland waters. but are still much greater than in the ocean. the These changes are not so great as in the atmos- phere. even in the is which polar regions. Seasonal variations. Such conditions again naturally at bring about a definite selection of fauna. shallower the water. as well as daily variations. are very great. aided by winds this overturn continues until the whole mass is cooled to 4. The lake trout ascends only the former for spawning. The peculiar temperature stratification in lakes resulting from the sinking of the cooled surface waters will be discussed in more detail in Chapter 18.000 to 40. other things being equal. so that there are great variations in temperature. have heat is ' budgets (of the water itself but not of the lake as a whole) on the order of 30.352 Environmental Factors in Inland Waters has two tributaries. The confined to the surface by the fact that it is less dense than water. one of which is much richer in oxygen than the other. is rendered extremely variable by their small size. These lakes of approximately the latitude of Chicago tend to have a somewhat higher and more uniform heat budget than . pure water is densest 4. forms. Unlike sea water. The shallower depth of many allows a rapid and intensive heating during the day but also an equally rapid cooling at night. It is because of these facts that water a few meters deep. Finally a covering of ice colder strata of water are J . like their chemical content. in length by 2 or more km. and in extreme cases the waters may evaporate or freeze. Heat budgets of lakes The amount of heat necessary to melt the ice and to raise the temperature of the water of a lake from the winter minimum to the sum- mer maximum 1 10 called the annual heat budget.

Multiplicity of Niches

353

those of Europe, and there is no evidence of an increase in the annual heat budget from latitude 40 to 60 N. For most lakes, the majority 2 10 of the annual heat budget is distributed as a result of wind action.
'

Light penetration
the ocean, many factors influence the penetration of light into fresh waters; of these the amount of suspended matter is particularly effective in excluding light. Crystal Lake, Wisconsin, with highly
in

As

transparent water, has approximately the same light penetration as Puget Sound. In Wisconsin lakes, the depth at which light is re-

duced

to 1% of that at the surface

was found by Birge and Juday

to

2 vary from 1.5 to 29 m.

The amount

their shallowness.

of light in inland waters is markedly influenced by Only a few of these waters are more than 300-

400 m. deep; most of them are shallower than 30 or 40 m.
in the

But even

deeper ones, the shallow shore region, despite its smaller surface far better populated than the deeper water. This comparison is area, is the more striking when considered in connection with similar reLight penetrates to the very bottom of many lakes rich flora rooted in the lake floor. The roots of plants make use of the mineral substances of the bottom much better than if these were merely dissolved by the standing waters above
lations in the sea.

and makes possible a

them; and the production of organic substances is accordingly increased. Besides this, vascular plants thrive in much greater abun-

water than in the seas; these also are softer and are more easily accessible than land plants as food for snails and insect larvae, because they need no protection against diying and less me-

dance

in fresh

chanical supporting tissues; they therefore disintegrate faster after dying off, and furnish food for the detritus feeders. For these reasons

the fresh waters in general are much more thickly populated with Lohmann, in comparing the value of living forms than the ocean.

the catches in the nannoplankton of the tropical oceans, of the cool open ocean, of coastal waters of the Bay of Kiel, and in fresh water,

A lake in Holstein produced 217.5 cc. got this ratio: 1:10:988:9017. of plankton per cubic meter of water, while in the ocean the whole mass of water under 1 sq. m. of surface area produced only 150180 gm. of organic substance.
Multiplicity of niches The multiplicity of the environmental conditions in inland waters
great.

is

Every gradation

in

water movement

is

found, from foaming,

354

Environmental Factors in Inland Waters
tumbling from rock to rock in the mountain torrents,

falling water,

The standing waters, to the lazily flowing streams of the plains. especially those of larger surface area, are often whipped into waves by the wind and beat heavily against their shores, while many ponds
have placid surfaces and stagnant depths. Tides are practically absent; about 5-cm. tidal differences have been reported for Lake Michigan. Their place is taken to some extent by the seiches,* which also occur along seacoasts. Just as in the ocean, water movement has a There are rheocolous animals, decisive influence on living forms. adapted to a more or less strong current, and limnocolous, which
thrive only in quiet waters.

The manifold gradations

in

chemical content, temperature,

light,

and water currents are again variously combined in the separate bodies of water. There can be no doubt that the far-reaching variation of living conditions in inland waters is primarily a result of the small size of the bodies of water as compared with the ocean. The

numberless possibilities of special differentiation under the influence of the environment determine an immense wealth of species of animals, and this is further increased by very decided isolation. Standing waters are often entirely isolated from each other and, with their tributaries, often form separate systems whose contained life is con-

nected only by passive or accidental distribution. Running waters, to be sure, are continuous over greater areas, but the individual river And even systems are separated from each other by watersheds.

though they all empty into the ocean, this forms an insuperable barrier for most forms because of its salt content. The wealth of species in the inland waters differs from that of the ocean. In the ocean there are much more numerous structural plans within which the species are invariable over wide ranges. In inland waters the number of types is limited, but within individual species an almost unlimited variation is the rule, at least among the less vagile forms, so that one might almost say, as the number of lakes, so the

number

of geographic forms. Variability of animals in inland waters, especially in fresh waters, may be further illustrated.

three

Ceratium hirundinella (Fig. 80), one of the armored flagellates, has and four horned forms, and the size varies between the extremes 13 found that of 92 n in Lake Como and 707 p, in Lake Schwendi. List had its local is definite which form characterized by the every pond

*A
varies

seiche (sache) is an oscillation of water above and below mean level; from a few centimeters to 1.5 m. and is usually attributed to variation

it

in

atmospheric pressure or to the action of a strong wind.

Multiplicity of Niches
relative lengths of the horns

355

and that there is a marked difference according to whether Ceratium is found in shallow ( 2-4 m. ) or deep (4-16 m. ) ponds; he proved by experiment that this depends on the direct influence of the environment. The snails and mussels vary in Germany from pond to pond and from one river system to another; similarly the lakes in Celebes have their own local forms of Melaniidae and Corbiculidae, 12 and each of the Patagonian river systems, which

FIG. 80.

Switzerland:

Ceratium hirundinella, small and large forms from the same lakes in 1, from the Untcrsce; 2, from Lake Lucerne; 3, from Lake MagAfter Bachmann. giore; 4, from Lake Como.

are especially effectively isolated, contains its own groups of mollusks that are distinctly different from those in adjacent rivers. 17 The lower Crustacea give excellent examples of the endless variation of species in inland waters. The brine shrimp Artemia salina varies so greatly that almost every salt lake has its own race. 19

Numerous
by

this great variation.

species of Cladocera of different genera are distinguished Among Bosmina coregoni the end forms are

distinguished

by a

large

number

of characteristics but are united
localities.
18

by

unbroken

In the genus Daphnia twenty-eight forms, once described as different species, are now included in the species D. longispina; others are similarly vari22

transitional chains

from various

reports similar local variations for the African freshwater crab Telphusa perlata. Just as the whitefish Coregonus disable.

Weber

23

plays peculiar characteristics from lake to lake, so the European

356

Environmental Factors
trout

in

Inland Waters

and the European lake trout S. lacustris also vary considerably, and these two are so closely connected by transi4 The tional forms that they have even been placed in one species. American black-spotted trout (west of the Great Plains) also have 9 In Australia almost every innumerable subspecies and local forms. 14 Such examples river has its own variety of the salmonoid Galaxias. be fishes might greatly multiplied. among

brown

Salmo

fario

Although many inhabitants of the inland waters are subject to mutation and show exceptionally great variation within the species, the fauna of inland waters the world over maintains a marked uniformity. Species of world-wide distribution occur everywhere, along with regionally distinct species and genera of limited distribution; these are sometimes called universal, as contrasted with the regional,
fresh-water fauna.
cies has impressed water fauna stands

The
in

surprisingly great
investigators,

number

many

and

of widespread spein this respect the fresh-

of the land.

sharp contrast with that of the ocean as well as The explanation suggested previously (p. 102 ff.) was

that the short life of a given habitat in the inland waters has conditioned the evolution of forms capable of ready transfer by accidental

or incidental emigration. Local forms incapable of such transfer normally perish as their habitats change. It is sufficient merely to call
attention to this matter here.

Fedonic and limnetic organisms In the fauna of inland waters as well as
water.
It is

in that of the

ocean one

can distinguish between animals of the bottom and those of the open
natural for these groups to retain the names benthic and pelagic, as they are designated in the ocean; but it has become a common practice to speak of the bottom organisms of the fresh water
as
in fresh

pedonic and pelagic forms as limnetic. Among the pedonic animals water there is also a distinction between shore forms and

animals found at greater depths; practically speaking, the deep-water community begins at the border of plant growth. This level varies with the transparency of the water and usually lies between 7 m. or
less and 30 m. A sunless deep stratum with a truly abyssal fauna occurs in only those few lakes whose depth is over 400 m., i.e., especially in Lake Baikal and Lake Tanganyika. The giant planarians and the non-pigmented fish (Comephorus baikalensis) of Lake Baikal are

truly abyssal fresh-water animals.

In the limnetic fauna of inland

waters,

we

again distinguish between the drifting plankton and the

Pedonic and Limnetic Organisms
nekton, which
is

357

independent and swims without the aid of water movements. The nekton includes only vertebrates in fresh waters. The pedonic fauna of fresh water differs from the corresponding fauna of the ocean in that sessile animals, which are so plentifully
represented in the benthal of the ocean, are almost entirely absent, with the exception of several ciliates, fresh-water sponges, the fresh-

water polyp Hydra, certain bryozoans, and a few attached insect larvae. Emigration of sessile animals into rivers against the current is' probably difficult. Except certain bacteria and an aquatic glowworm, no fresh-water luminous organisms are known. 5 The limnetic fauna, especially the fresh-water plankton, has, in common with that
of the ocean, important adaptations that facilitate floating. Both the limnetic and the pedonic environments are poorer in number of species than are corresponding regions in the ocean, since so many types are

completely absent in fresh water, but, although it is qualitatively The almost complete abpoorer, quantitatively it is much richer.
very noticeable in the composition of freshwater plankton in comparison with the plankton of the ocean. Only the mussel, Dreissena polymorpha, which has been introduced into
is

sence of larvae and eggs

by navigation, has planktonic larvae; besides these there are the flagellated larvae of some Cestodes and the nauplii of many copepods. In open water, the statoblasts of many Bryozoa and the
winter eggs of many rotifers and of a few cladocerans are found; the property of adhesion to the surface of the water, which these small
bodies possess, gives them the advantage of being driven by the wind into the littoral region, where they find a higher temperature, which promotes development. 26 The glochidian larvae of the lake and

fresh water

(Anodonta, Unio) and the lake-floor larvae of midges Chironomidae Tendipedidae) are occasionally mixed with the ( of the wave action of shallow waters. means plankton by
river mussels

=

The character of the fresh-water plankton is further considerably influenced by the fact that the size of its animals is much less than in the ocean. Such large forms as siphonophores, ctenophores, heteropods, pteropods, and arrowworms are not known in fresh waters.

The marine animals

just

mentioned are mostly

jelly-like; their

bodies

contain an extraordinary amount of water, which considerably increases their size. The absorption of large amounts of fresh water into

however, impossible without damage to We know of jelly-like matter in only a the organism (cf. p. f.). few cases in fresh-water animal plankters and in these only lifeless the mantle of the water flea parts of the body are expanded, e.g.,
the

body

of an animal

is,

43

358

Environmental Factors

in

Inland Waters

Holopedium

The increase (Fig. 81) and a few rotifers (Fig. 82). of surface area by means of thread-like pseudopodia, such as serve

as aids to floating in Radiolaria and Foraminifera in the ocean, though found in the Heliozoa, occurs much more rarely among fresh-

water animals, perhaps because with such greatly increased sur-

FIG. 81.

Holopedium gibberum.

X 14.

After Fri

and Vavra.

face area the

amount

of fresh water absorbed

would be too

great.

Non-gelatinous plankton animals also reach a greater size in the ocean than in fresh water, which may be attributed to the lesser density and 2* consequent lessened buoyancy of fresh as compared with sea water.

The The

skeletons of fresh-water plankton animals rarely contain lime. largest animal of fresh-water plankton, the larvae of Corethra plumicornis (Fig. 104), which reaches a length of 15 mm., has two

pairs of air-filled tracheal bladders that enable it to float. The minimum size, however, is about the same for the fresh-water plankton

as for that of the ocean; there is a dwarf plankton in both regions. Secondary inhabitants of this medium, those which have returned

Pedonic and Limnetic Organisms

359

from the terrestrial to the aquatic habitat, are as plentiful in fresh water as they are scarce in the ocean. They include snails, insects, The varied conditions of water movearachnids, and vertebrates. ment and the necessity for getting oxygen have brought about many
convergent structures,
beetles (Dytiscidae
e.g., the flat, sharp-edged body shape of water and Hydrophilidae ) and water bugs (Naucori-

FIG.

82.
in

Mastigocerca
its

sett-

fera

X

130.

gelatinous case. After Lauterborn.

Heads of a frog, a crocoand a hippotamus as examples of convergent adaptation to amphibFIG. 83.
dile,

ian

life.

of the

dae); the swimming legs, widened by means of hairs; the lightening body by the carrying of air on the hairy abdominal surfaces,

beetles (Hydrophilus) and bugs (Notonecta); and the development of long breathing tubes that reach to the surface, among ani-

among

mals of shallow waters,
the rat-tailed

e.g.,

some water bugs (Nepa, Ranatra) and

maggot of the hover fly (Eristalis). Many vertebrates are amphibious; they avoid heat, evaporation, or enemies, or some combination of these, and often find their hunting grounds in the

They live periodically out of water; some of these have a strikin that they can immerse their bodies in the water up similarity ing to the nostrils and eyes (Fig. 83). The soft-shelled turtles (Trionywater.

360

Environmental Factors

in

Inland Waters

chidae) have their nostrils situated at the end of an elongate snout, so that they compare rather with Nepa and Ranatra (with breathing tubes at the rear end of the body) than with other aquatic reptiles.

One

fresh waters

of the primary distinctions between the animal communities of and those of the sea lies in the great proportion of such

terrestrial invaders of the inland waters.

BIBLIOGRAPHY
1.

2.

"Heat Budgets of American and European Lakes." Trans. 18:166-213, 3 figs. Birge, E. A., and Chancey Juday, 1914, "The inland lakes of Wisconsin. The hydrography and morphometry of the lakes." Bull. Wise. Geol. Natural Hist. Survey, 27:xv, 137, 8 figs., 29 maps.
Birge, E. A., 1915,
Set.,

Wise. Acad.

3.

Germain,
2:322.

L.,

1909, "Hydrobiologische Referate."

Intern.

Rev. HydrobioL,

4. Goldi,

E. A., 1914, Die Tierwelt der Schweiz in der Gegenwart
I.

und
1

in

der
(p.

Vergangenheit.

Wirbeltiere, Bern, Franke:

xvi,

654, 5

pis.,

map

578
5.

ff.).

Harvey, E. N., 1940, Living
1 pi.,

light.

Princeton, Princeton Univ. Press: xv, 328,
Jena, Fischer:

88

text figs.

6.

Hesse, Richard, Tiergeographie auf oekologischer Grundlage.
xii,

613, 135

figs.

7.

Hubbs, C.

L., 1940, "Fishes of the desert." E., 1935,

The

Biologist,

22:61-69, 7

figs.

8. Jewell,

Minna
S.,

"An

ecological study of the fresh-water sponges of
pis.

northeastern Wisconsin."
9.

Ecol Monographs, 5:461-506, 3

N. Y., Holt, 2 1905, Guide to the study of fishes. 10. Juday, Chancey, 1940, "The annual heat budget of an inland lake.*
Jordan, D.

vols., illus.

Ecology.

21:438-450.
11. 12. Kruimel, J. H., 1913, "Verzeichnis

Korwenkontio, V. A., 1911, "Einzelreferate." Intern. Rev. Hydrobiol, 4:521. der von Herrn E. C. Abendanon in Celebes

13. List,

gesammelten Susswassermollusken." Bijdr. Dierkunde, 19:217-235. Theodor, 1913, "Uber die Temporal- und Localvariationen von Ceratium hirundinella O. F. M. aus dem Plankton einiger Teiche in der Umgegend von Darmstadt und einiger Kalke des Altrheins bei Erfelden." Arch. Hydrobiol.,
fig.

9: 81-126, 1
14.

Lucas, A. H. S., 1897, "On some facts in the geographical distribution of land and fresh-water vertebrates in Victoria." Proc. Roy. Soc. Victoria, (n.s. )

9:34-53.
15. Morrison,
J.

E.

P.,

consin lake districts."
16.

1932, "A report on the Mollusca of the northeastern WisTrans. Wise. Acad. Set., 27:359-396, 127 figs.
J.

Mrazek,

Al.,

1903, "Ergebnisse einer von Dr. Al. Mrazek im

1902 nach

17.

Montenegro unternommenen Sammelreise: I. Einleitung und Reisebericht." Sitzber. bohm. Ges. Wiss. f 1903, No. 15: 1-24, 4 pis. Rept. Princeton Pilsbry, H. A., 1911, "Non-marine Mollusca of Patagonia." Univ. Exped. Patagonia, 3:513-633, 20 pis., 38 text figs.

Bibliography
18.

361

Ruhe, F. E., 1912, "Monographic der Daphniden Deutschlands und der benachbarten Gebiete: I. Monographic des Genus Bosmina. A. Bosmina coregoni im baltischen Seengebiete." Zoologica (Stuttgart), 25: Heft, 63:1141, 7 pis., 1 text fig., 3 maps. 19. Samter, M., and R. Heymons, 1902, "Die Variationen bei Artemia salina Leach und ihre Abhiingigkeit von ausseren Einfliissen." Abhandl. Akad.
Wiss. Berlin, 1902, Phys. Abhandl, 2:1-62.
20. Stephens, Jane, 1912, Porifera."

"A biological survey of Clare Island: 58. Fresh-water Proc. Irish Acad., 31, Sect. 3, No. 60:1-17, 1 pi.

21. Volk, Richard, 1907, "Mitteilungen iiber die Biologische Elbe Untersuchung des naturhistorischen Museums in Hamburg." Verhandl. naturw. Ver. Ham-

burg, (3) 15:1-54, 3

pis.,

1

map

(p. 29).

22. Wagler, Erich, 1912, "Faunistische

und

biologische Studien an freischwimpis.,

menden Cladoceren
14 text
23.
figs.

Sachsens."

Zoologica (Stuttgart), 26:306-366, 30

Weber, Max, 1897, "Beitrage zur Kenntnis der Fauna von Siid-Afrika. Ergebnisse einer Reise von Prof. Max Weber im Jahre 1894: I. Zur Kenntnis der Zool Jahrb. Syst., 10:135-200, 15 pis. Siisswiisser-Fauna von Siid-Afrika."
(p.

156).
P. S., 1935,

24.

Welch,
168).

Limnology.

N.

Y.,

McGraw-Hill:

xiv,

471, 46

figs.

(p.

25.

1909, "Ober die praktische Bedeutung der jahrlichen Variationen in die Viskositat des Wassers." Intern. Rev. Hydrobiol., 2:231-

Wesenberg-Lund,
233.

26. Idom,

1909, "Uber pelagische Eier, Dauerzustande und Larvenstadien der pclagischen Region des Siisswassers." Ibid., 2:424-448, 3 figs.

-//

Communities

in

Running Waters

INLAND WATERS MAY BE CLASSIFIED PRIMARILY INTO FLOWING, OR LOTIC, and standing, or lentic, waters. It is desirable to separate the standing waters into the fresh-water lakes and ponds proper and, on the other hand, into the salt lakes or pools, which contain in solution large amounts of such substances as sodium chloride, magnesium sulphate, and humus. It is also useful to give separate treatment to subterranean waters, whether found as ground water or in caves or
wells.

There are certain regularly recurring differences between the inhabitants of running and standing waters that necessitate a separation of the two. Differences in area, movements of the water, and the differing relationships of depths and of temperatures bring about divergences, which may become extreme. These two types or water environment are united by very gradual transitions- a river of th*> plains in which the current is hardly noticeable (as in many of the steppe rivers of south Russia in summer), a lake through which a river flows, and an ox-bow lake through which there is a current only
during floods offer examples of situations in which a difference between the faunae of running and standing waters is hardly perceptible. Furthermore, the animal communities of the wave-washed rocky eroding shores of lakes are closely similar to those of neighboring rocky rapids in streams.

Departing from common custom, we shall consider the running waters first because of their relatively long geological duration, their

more extensive

continuity, and their usual connection with the ocean. furnish the routes for the active dispersal of living forms from They the ocean into fresh water, which is still going on. Fishes, snails,

mussels, and larger crustaceans, which are capable of considerable movement, have thus reached rivers and lakes, as have less active ani362

River Plankton

383

mals such as leeches and other annelids, turbellarians, and the lower
crustaceans.

The composition
lakes, for instance,

makes

of the deep-water communities of the lower Alpine it very probable that at least a part of their

came in at the end of the glacial period by way of the streams large flowing from the melting glaciers. Running waters thus formed a highway by which arctic animals moved into the lakes of
inhabitants

the Alpine foothills, and perhaps, vice versa, mals reached the Scandinavian lakes.

some
less

of the Alpine ani-

Chemical differences among river waters are

than those

among

standing waters. The mingling of waters from the various parts of a more or less large, often also geologically varied, river basin equalizes

chemical variation

much more than

in standing waters.

Even

when
by

very salty or boggy streams enter a river, the water contributed them is as a rule negligible in comparison with that of the main

There is great difference in turbidity in streams, from the clear water of mountain brooks to the perpetual muddiness of such rivers as the Missouri. At flood, many streams, otherwise clear, become mud-laden torrents. The whole phenomenon of flooding, with
stream.

accompanying frequently drastic changes in level, strength of current, width of stream, etc., makes another difference between lentic and lotic environments. Rivers, furthermore, usually have no very distinct deep stratum; and even in such large rivers as the Mississippi, the Amazon, and the Congo, where the depth is great, we do not know of any special deepwater communities.
River plankton

have no plankton of their own in the sense animals or plankton communities found only in of typical plankton rivers. There is, of course, a floating fauna in rivers; this is not, how-

Running waters,

finally,

ever, autochthonic but has

its

origin in standing waters, in lakes

through which the rivers flow, in ox-bow lakes connected with the The river plankton is inrivers, and in quiet bays and back waters. creased by floods that flush out such places, and thus species are added to the river plankton that do not otherwise occur there; in the plankton of some rivers of south Russia, the larvae of the branchiopod 24 Limnetis were found after a flood. They ordinarily occur only in
the transient water basins of the steppe. River plankton, consequently, is sparser than that of the lakes from which it originates. Investigations at the outlet of

Lunzer Lake indicate that the stronger swimming

364

Communities
e.g.,

in

Running Waters

plankton organisms,

crustaceans and even rotifers, tend to

move

against the current in the region of outflow and, unless the current is swift, such forms are thus enabled to move into the river in relatively

small numbers. 32

The plankton communities

of different rivers,

how-

ever, are distinct, as are those of the lakes through which they flow. So the plankton of the upper Rhine below Lake Constance differs from

Aar as the plankton of Lake Constance differs from that of Zurich, and the plankton of the Neva is almost identical with that of Lake Ladoga, of which it is the outlet, whereas its companion river, the Tosna, contains a wholly different plankton, which is explained by its bog origin. Evidence from a different approach supports the generalization that dissimilarity of habitat is more effective
that of the

Lake

than separateness in determining distinctiveness of plankton populations. Two small lakes near Lake Huron have almost identical plankton and contribute about equally in kind, annual maxima, and mass to the population of the river. This likeness probably depends on the
similarity of substrate and of drainage basins of the two closely neigh1 boring lakes. Speed of current and short stream length are apparently the main causes for rivers not developing their own plankton. The Neva, which extends for 60 km. from its source in Lake Ladoga to the ocean, flows

m. a second; a mass of water with its plankton, therefore, reaches salt water in 12 to 14 hours; this time is too short for any considerable multiplication, even among animals that reproduce rapidly. The water in the upper Rhine, flowing at a rate of about 2.5 m. a second, covers the distance between Basel and Manitheim in 1% days. The conditions become more favorable with the reduction in their gradient, which occurs in most rivers near their mouths, and in some much sooner. The river then tends to develop
at a rate of 1.0

m. to

1.7

and maintain its own plankton. The waters of the Moskva cov^r only between 14 and 30 km. in 24 hours; consequently, short-lived animals that reproduce rapidly can greatly increase their numbers on their way to the ocean rotifers, for example, whose numbers tend to be

much

greater than those of Crustacea in slowly flowing rivers. For the same reasons there is seldom a monotonous plankton in a river, such as would result from mass development of any one species.

In addition to the effect of their quickened run-off, rapids often serve to destroy plankton organisms. In the Mississippi River, the

amount

of plankton immediately below Rock Island Rapids is less than half that just above. The passage of these rapids requires about 8 hours; the cause of death is thought to be the violent impacts against

Stream Subdivisions
10 suspended sand grains or against the bottom.

365
Mats of submerged
as

vegetation also

remove plankton; a decrease of
in

much

as 503! has

been observed within 20 m.

dense aquatic vegetation. 80 Hydropsyche nets, if present in numbers, take a heavy toll of passing plank20 ters, especially in shallow rapids.

A
tide

peculiarity of river

mouths

is

the piling

up of the water
this,

at high

and

its

mixture with

salt water.

Because of

the outflow of

otherwise slowly moving rivers is temporarily stopped, and the conditions resemble those in standing waters. The remarkable richness of
the river plankton in such regions is no doubt connected with this fact. Thus a cubic meter of water in the Elbe above Hamburg contains a

few thousand cladocerans, but below Hamburg the number

risos to

many
mina

millions, in the India Harbor, for example, to 11,040,000 BosMost important of all, forms adapted to brackish longirostris.

water become mingled with the water of the river mouth, and some of these increase to great numbers. The copepod Popella guerni is found
in the Volga delta; and in the mouth of the Amazon the copepods Weismanella and Pseudodiaptomus gracilis occur. These are marine

components of

river plankton.

Stream subdivisions

The
as
lar
it

divisions of a flowing stream according to the amount of water, diminishes from mouth to source, which correspond to the popuriver, creek, brook, rivulet, spring,
its

names

have

little

to

do with the

composition of
of erosion

fauna.

Division into lower river (with a

minimum

and a maximum of deposit), middle river (with a balance between erosion and deposit and a more noticeable lateral erosion), and upper river ( with a maximum of deep erosion and a minimum of
also frequently inapplicable. An important factor for anithe velocity of the current, and, in correlation with this, the nature of the substratum, the temperature of the water, and its supply But the fall of a river does not increase steadily from of oxygen.

deposit)

is

mal

life is

mouth to the source. The course of the Rhine, for instance, has three regions of very rapid current, between which there are regions of much more gradual fall. The Amazon is a stream of minimum fall
the

through most of its course; Manaus, 1400 km. from the mouth, lies only 26 m. above sea level; the average fall in this region is 19 m. per 1000 km. In the great rapids in mid-course of many large rivers, such as the Congo or the Essequibo, there are animals with adaptive features

much

like those of

the inhabitants of small mountain streams.
river of the Skava, a Galician

The muddy Spytkowianka, a source

366

Communities

in

Running Waters

tributary of the Vistula, contains fishes that are otherwise characteristic of the lower courses of rivers, and only farther downstream is the fall
of the Skava sufficient for trout

and

grayling.

gradient mainly determines the physical nature of the substratum in streams. Transportable parts of the substratum are car-

The

and mud can be deposited only in protected quiet the bottoms of rapidly running rivers are covered otherwise coves; with coarse rock, the smaller pieces of which are constantly in motion. The slower the stream, the finer the fragments carried, until finally only very fine particles can be moved and only fine gravel is deposited;
ried along; sand
this
is

succeeded by sand and
is

lastly

by

mud

bottom.

As the organic

lightest and consequently the last to sink to the bottom, of nutritive sediment from the whole river basin collect in quantities

detritus

the regions of the slowest current, usually near the mouth of a river. This is a determining factor for the bottom fauna. All life is crushed
rocks; only on permanent cliffs, on especially large blocks, or in sheltered places, can animals gain a foothold in a moving water-rock environment.

among

constantly

moving

The greatest amount of life in streams with rock bottoms se^ms to be found on or among mixed rubble rather than in association with 160 Moving ice in spring and the ice rubble large or with small stones. that forms in falls and rapids constitute an additional destructive factor in northern and alpine waters. 16 Fine sand and mud, on the other hand, furnish an opportunity for burrowing and tunneling, and a
rich food supply for detritus feeders. The supply of oxygen is especially favorable in swiftly flowing streams with strong rapids and falls because of the immense water

surface that takes

up oxygen, and because of the thorough mixing. Even where the current is slow, the water is much better aerated

than in standing waters containing a similarly small popi ifotion of

green plants.

The

daily

and yearly

fluctuations of temperature are

much

smaller

in rapidly flowing mountain streams, often than in the waters of the plains, which are

shaded by the mountains, exposed to sunshine. The fluctuations are least in the springs that emerge from deep underground. The amount of the annual fluctuation of temperature in streams from glaciers is to 1; in high mountain springs, 1 to in in barbel rivers 6; lower parts of trout brooks, 15 or more; perhaps about 19; in carp rivers, about 24V Even in smaller streams with little movement, the temperature of the water is more uniform than that
of the
air.

Stenothermal cold-limited animals, as well as eurythermal

High tide has no It is characterized by weak swimmers with effect on this region. Scardinius. After Bloch. with a partly soft and partly gravelly substratum. swiftly running water that is not entirely clear. compressed bodies (Fig. with similar characteristics though without a mixture of sea water. ing too many points of vantage for the current. and deep. and the miller's thumb. bream. eryth rophthalmus. moderately warm. of standing waters. 367 more or less euryther- fall of a stream and its accompanying environmental factors a sorting out of the fauna. Next upstream is the barbel region with deep. the current weak. Cyprinus carpio. live in rapidly flowing streams. . Misgurnus. but only mal animals can live in streams of slow current. The characteristic fishes are smelt and stickleback. inlaterally cluding the eelpout. is next upstream. In general this is such that the steeper gradients demand more distinct adaptations and thus contain fewer species. bitterling. the carp. 84. besides flounder and sturgeon in the lower regions and ruff (Acerina) and face. Abramis. Cross sections of fishes of swift streams (upper) compared with those Left to right. Carassius. European minnow. and Carassius vulgaris. Cottus gobio. bottom is soft. roach. 84). together with Leuciscus leudscus and L. and Rhodeus amarus.Stream Subdivisions animals. carp. lower. crucian carp. Cottus. since their muscles are weak and their bodies flat. mainly different from those of slack water. In the The effect river mouth region of streams of central Europe 12 the water is al- ways brackish near the bottom and at least occasionally near the sur- Fic. Phoxinus. upper: trout. miller's thumb. red-eye. the eel in the upper.* Besides these offer they there are many fishes from the upper region of stronger currents. RJwdeus. the water muddy. Lota lota. Cyprinus carpio. unsuited for swimming in movwater because. warm. * The extremely flattened fishes are want- Chief among these are Abramis brama. Salmo fario. The carp region.

miller's-thumb). finally. the barbel. there is an 23 essentially similar distribution of fishes in the Chicago region. The boundaries of these regions naturally are nowhere sharply defined. With an increased fall the number of accompanying fishes decreases more and more.* After considerable. when 41 species of fishes were known Li Holland. Wherever the classification can generally be applied. Graylings. between 1500 and 2100 m.. above sea level. region. the charA acteristic fish. after Lake Constance. Phoxinus laevis. (trfrpt. deep. this gives a clear illustration of selection through severe factors of environment. tench. only 13 species go beyond 2750.. 11 they included 22 Cyprinidae. there are 44. up are the minnow. while its upper course may correspond to the carp region and may contain carp. In the Rhine. The distribution of fishes according to height above sea level in Colorado is similar. and 9 migrants f oni the sea. occur. In the upper Rhine. Cobitis barand the miller's-thumb. absent in the carp region. such as minnows of the genus Leuciscus. or trout. min8 now. are also found as low as 2100. includes brooks and smaller rivers with rocky and coarsely graveled bottoms.. moderately Communities in Running Waters compressed forms like the roach still occur. their order may even be changed according to the topography of the bottom. search. cool water with stronger currents. for instance. and above the to falls 28. * . there were 33 species. as a rule.368 ing. but may be absent. Besides grayling there are a number of fishes from both neighboring regions. with sandy or partly soft substratum. Almost all those that occur above 2750 m. and pike. The characteristic form is the trout Salmo fario. All the fishes of this region have rounded bodies (Fig. 11 species went as high as 700 m. The trout region. those going farthest batula. 47. and only 3 above 1900 in. grayling region usually forms the transition to the uppermost. 84). below the falls. either for the Rhine or for the streams of Colorado. 5 as high as 1100 m. and this number decreases to 24 between 2100 and 2750 m. has a rounded body. 7 below 1500 m. only the subtraction of 3 whitefish (Corcgonus) confined 25. Though the species differ. we are unable to find a more modern statement of these interesting distributional relations of fishes. 4 salmon. and the non-salmonids are found in the still lower regions. 7 of which are salmonids. Barbus fluviatilis. moderately The warm. and clear. with rapidly moving water. there is a decrease in number of species of fish from the mouth to the source of the river. It includes larger creeks and middlesized rivers. the roach. limited number of stenothermal forms also occur. and a creek may be typically a trout region in the middle of its course.

For great distances in the upper Alps and the highlands north of the Alps. that it is a relic of the glacial period and was found in all the streams of middle Europe during that time. After Voigt. 23 ' 33 animals other than fishes are restricted by velocity of current. and PL . lives in warmer waters. and its distribution indicates ing. and PL alpina is often limited entirely to the source brooks or perhaps springs at the actual source. The Pohjcelis cornuta. The stratifying influence of temperature is especially marked FIG. among European brook planarians. Poly cells cornuta. Planaria gonocephala. but optimum temperature is several degrees higher. are distributed so that the first lives in the warmer waters with stony bottoms (Fig. The brook planarians of central Europe. P. 85. In the warmer springs of the lower regions PL alpina and Pohjcelis are entirely missPL alpina breeds in the winter. so that the present distribution of fishes in a stream with varied types of environment from mouth to source gives us an insight into the general phenomena of ecological succession in streams. 85). on the other hand. mudbottom type as an end stage. Pohjcelis further up. substratum. Planaria gonocephala. 86).Stream Subdivisions 369 The physiographic history of many streams includes a transition from the swift-water and rock-bottom stage to the slow-moving. characteristic of running water in three species of flatworms Germany. In the middle region of the highlands the regions of distribution of the three species are rather closely ap- proximated (Fig. Pohjcelis cornuta also breeds in the winter. and PL alpina. although it can tolerate low temperatures. cornuta moved into the creeks after PL alpina. while PL alpina lives in the upper streams and springs. and temperature to certain definite regions in flowing Many waters. Top to bottom: Planaria alpina. PL gonocephala. PL alpina is the sole occupant. its optimum temperature its is below 10.

Polycelis is eliminated by the two Planaria. and Planaria gonocephala (V) succeed each other This with areas of intermixture (II and IV) between the adjacent species. section of a original arrangement is is altered when the conditions are changed. 86. Polycelis cornuta (III). Planaria alpina (I). replaced by Polycelis. under primitive conditions. in B. in C. In A.370 Communities in Running Waters ^iSSpiKSiSSS^fi Distribution of brook planarians shown on a schematic longitudinal mountain stream. Planaria alpina After Voigt. FIG. .

especially the small Sphaerium and Pisidium and the much larger Unio and Anodonta. Piece of clay from the bank of the Marne. 87.1 belong to the detritus feeders. Longitudinal section through one of these burrows. others with single openings. below the water level. The banks of many slowly flowing streams are riddled by the ramifying burrows of Ephemeridae^Fig. with substrata of mud or fine sand and with slow currents. which have their anends stuck in the mud from which they extract their food. riddled with the burrows of mayfly larvae ( Palingenfa ) . Bryozoa also storm. some with double. are especially plentiful. the terior alderfly Sialis. since each species is most efficient in its own optimum temperature. . 2 ' 1 "-. either creeping along in the surface layer (Sialis) or building mud tubes (Chironomus) or digging deeper bur- rows. They usually burrow in the mud. and several mayflies. also in other river regions with similar conditions. b. the midge Chironomus. After de Reaumur. Tubificid annelids. Many insect larvae also feed on detritus.. Their distribution in the brooks is determined by their competition for nourishment correlated with temperature. like those of muddy floors of lakes and ponds. These are plentiful in the Elbe and . they are found in such numbers that when a mass of subimagoes emerges at the same time on a sultry evening the air is filled with them as with a heavy snow- Mussels also occur. the inhabitants of the bottom are to a great extent detritus feeders. In the Elbe below Hamburg. without coarse sediment in suspension. e. The carp region In the lower courses of rivers. and FIG.g. 87). Sphaerium rivals the Tubificidae in numbers.The Carp Region 371 gonocephala followed still later.

These include principally species coming up to spawn. Amazon. otters and river dolphins (Platanista in the rivers of India. but need a firmer foundation var. Crocodiles and river turtles. reaches of the river. Cristatella. and suckers. rivers. and many others. the African Mormyridae. Experience in fishing shows water he must have. water the the leech Asellus. the farther a fisher- detritus feeders. and many rays have entirely adapted themselves to life in the The abundance of nourishment in slow streams. There are the flatworm Procotyla Valvata. water. Increased velocity of current and prevailing gravelly and stony bot- tom. also attracts a in the number of higher vertebrates to these regions. Fredericella. Paludicella. The flounder (Pleuronectes platessa) comes up into European rivers only in search of food. and Bythinia are rather plentiful.. Thus Plumatella princeps snail shells until they large development of lower animal life on the nutritive elements of the thick layer of mud and detritus on the bottoms of slowly flowing streams affords the basic food supply for the large numbers of fishes found in the lower regions of streams. spongiosa overruns resemble small potatoes. * they do not. Herpobdella. catfishes. and Into in South America) feed on the plentiful supply of Sirenia etc. e. together with sturgeons and spoonbills. yearly net profit per hectare (about 2. especially tropics. and it never becomes sexually mature in fresh. the gill-breathing . and occur in a number of beautiful forms. As a matter of lives river. Other invertebrates are somewhat less abundant than the detritus feeders. 14 Fresh-water sponges (Ephydatia fluvialitis) attach themselves in like manner. live on the mud. Pectinatella. the greater the area of man from the mouth of a course the number of migratory fishes is also largest in the lower Many stop here. predaceous fishes in turn are associated with the herbivores and that.! for attachment. and others. Theodostts. Among snails. These are primarily The bottom feeders such as carp. isopod fluviatilis. forms such as Viviparus fasciatus. and those that go farther upstream must also pass through the lower river. however.g. conditions generally found * t the middle courses of Lophopus. Salmonidac and sturgeons. from the ocean and feed there on water plants. at least at high water. (Manatus and Halicore) come up the rivers (Congo.5 acres) from fishing in extraordinarily good German lakes is about one-third that in the lower Oder and one-fifth that in the lower Elbe.372 Communities in Running Waters the Bille near Hamburg. In the tropics a number of selachians also enter fresh waters. which is subject to motion and consequent in friction. ) fish.

in bays or on the downstream side of gravel only banks. and living forms can find permanent attachment only on the firm rocks and boulders that resist the force of the current. reduce the whole group of mud inhabitants. but the species are different from those found on the muddy bottom of the lower river.. The sparse. becomes noticeable in the upper streams. and trout brooks in their upper regions. a number of libellulid dragonfly nymphs. or brooks. river are Some of the fishes found in this part of a more characteristic of the barbel or the grayling region. for fist by tossing and rubbing. where the fall is less marked. are to be found in such places. other species of Ephemeridae. where the fall The mechanical erosion of the current with is its considerably greater. especially of fishes. of course. plants and animals are crushed and ground. usually undermining the bordering cliffs. as are also some small water beetles. of the gill-breathing snails. the settling of living forms on the bottom is very limited.r>f28 29 There are. stones the size of a or a head are easily carried along. there we find tubificid worms and river mussels. Salmon overcome the barrier interposed by falls by leaping and are thus able inaccessible to other fishes. Free-swimming forms remain only in quiet places in secluded pools. Insect larvae continue to be important. Instead of midge larvae and the burrowing Ephemeridae. the larvae of stoneflies (Perlidae). various gradations even here. to range into the back waters of streams Many mountain fishes in the Andes and Himalayas have an adhesive organ by means of which they climb . The whole bottom is stony.The Trout Region 373 streams. They occur in quiet places. and even in the trout brooks. The few freshwater. the The much more ' brooks of the intermediate mountains can be divided into salmon brooks in their lower reaches. more and more of those occur that take shelter under stones. and. the coarseness of the rubble thus varies with the elevation. however. and above all a great number of caddis worms with their stone cases. Perpendicular waterfalls of considerable height block the passage of many animals. wild cataracts and rapids becomes most evident in the upper mountain streams. and is covered with loose rocks and stones that are being sifted by the force of the current. Because of this movement of rubble. rapidly moving plankton has already been mentioned. The trout region influence of increased velocity of current. Goniobasis and Viviparus. air-breathing snails living in such biotopes also cling to stones. as do Physa and Ancylus fluviatilis. the mountain rivers. which are completely missing in the mud.

and Diptera.u. are stationary at this level. i. aside from the small number of those that tion of quiet places. these may be present in tremendous numbers. The caddis worm Hydropsyche spins a veritable plankton net in i>wift water. the In comparison with similar conditions in the surf region of the number of permanent sessile forms is small. sucking disks. are attached. Planaria of snails dependent upon effective anchorage. forming a thick black blanket of writhing life on the brinks of waterfalls. Their In central Europe they include the triclad Hat. and the swimming hairs of the quiet-water mites are wanting on the legs of the swift. Many types have developed means of maintaining themselves in the current. outnumbering all others in species and individuals. Lota.. claws. ocean.water forms. large Perlidae.. The fresh- water sponge Ephtjdatia and the bryozoan Plumatella occasionally encrust the stones on the bottom of the brooks. the remaining inhabitants of mountain stream?. are settle among the sp. however. such as the egg cocoons of flatworms and leeches and the eggs of the water mites. . many finding refuge beneath the larger stones. besides a few Odonata and Perlidae are almost completely confined to rapidly flowing waters. the water flea (Gammarus pulex) and the river crayfish. the water mites have shorter legs than those in quiet waters. The bottom fauna is well adapted to mountain streams. a variety of water mites. rials Little larvae of the black nourishment is suspended in swift water. A few types of insects have larvae that attach themselves in swift water. mountain streams. whose relatives swim readily in quiet waters. Of the fishes.g. which will be discussed below. numbers of insect larvae of the groups Ephemeridae. the hairs on the swimmerets of the few ostracods are also much reduced. and Nemachilus hide under stones.je vegeta number is not large. stages that do not need nourishment.e. The Mountain brook adaptations The devices by which inhabitants of mountain streams are able to withstand the force of the current are unusually diverse. but often . Otherwise only the eggs or pupae.374 Communities in Running Waters Most swimmers. On the other hand. silk attachment threads. Other animals. and. e. Cottus. Coleoptera. Trichoptera. are excluded from vertical rock walls. or expanded pectoral fins.. The fly Simulium depend on the microscopic food mate- brought to their fan-like feeding apparatus by the passing current.g. alpina) and leeches (Glossiphonia] a number several small lymnaeids ) a f ew mus( Ancylus fluviatilis and sels (the brook pearl mussel Margaritifera margaritifera and species of Pisidium). worms (e.

g. while the larvae are attached by suckers or by a silken thread. Fig. 88. The flatworms and leeches. FIG. their cases weighted laterally by stones. in from swift streams. The body tends to be flat. After Bra nor. Odonata and Coleoptera. 89. the inhabitants of standing waters. shell of the snail Ancylus fluviatilis. such as Pscphenus.. 88). Silo. Shield- pendages. which have a flat circular "shell" extending well beyond the ap- Fie. FIG. After Ulmer. Most limpet-like of all are the larvae of certain beetles. larval Thremma. after arrow shows the direction of flow of the water. They are shaped larva may-fly from ( swift streams always small comparison with the larger cases of stoma). The legs of these larvae usually protrude laterally. 89&). The laterally compressed bodies of amphipods enable them to slip into narrow crevices. without marked protrusions. shield-shaped The in cases of caddis (e. and very many insect larvae. and the rim of the body is pressed closely to the substratum. 89 to FIG. at the same time 90) down . especially those of Perlidae and Ephemeridae (Fig. show this adaptation. a border of bristles often completes the attachment of the hard chitinous shield to the substratum. 90. water mites. Many pronounced rheocoles have developed a streamlined upper surface. and Silo nigricornis (b). worms may also be Thremma. The FIG. as a result of convergent adaptation. the upper surface smooth. the femurs are flattened. b. especially com- pressed at the head. the larvae fasten larger stones to their cases.Mountain Brook Adaptations 375 similar in forms of very different taxonomic relationships. A unique method of stabilization running waters is the weighting of the body. 89a). The shieldshaped pupa cases of the black flies ( Simulidae ) are X Prosopi6. 90 case of caddis fly Kobelt. Fig. often shield-shaped. after Klapalek. Larval cases of the caddisflies Gocra pilosa (a). Convergent adaptation swift water: a. the snail Ancylus (Fig. also attached to the substratum. In a few species of caddis worms (Gocra.

and the flat. from beneath. sole-like undersurface of the larvae of other midges of the families Blepharoceridae (Fig. 91. and the snails of this habitat have these surfaces much enlarged in comparison with land snails of equal size. to show their snrkleft. right. 910) and Psychodidae (Fig. 916) bears a longitudinal row of suction disks. a blepharocerid. a psychodid. Many mountain streams are adapted to life on the bottom by loss of the swim bladder. Organs of attachment of the most diverse kinds are especially characteristic of the all animals of this environment. Liponeura brevirostris. Peri- coma califarnica. surrounded by a capsule of bone. as in the Eurasian Cobitidae and the East Indian Homaloptera. ing disks: Dipterous larvae from swift streams. Fic. Flatworms and mayfly nymphs have strong adherent surfaces. After Wesenberg Lund. the posterior end of the larva of Melusina. since other fam6 ilies of salamanders also have a few lungless mountain brook forms. Many of the . The lungless mountain stream salamanders of North America ( Plethofishes of . after Brawr. bears sucking plates whose power of attachment is increased by bristles.376 Communities in Running Waters providing for a smooth attachment to the substratum by maintaining a level undersurface of these stones and by filling in the cracks. Leeches attach themselves by means of suction cups. as in Coitus and the darters ( Etheostominae ) or there may be only a small swim bladder. dontidae) apparently present a parallel adaptation. a biting gnat. The claws of the legs of water mites and insect larvae are strongly developed.

Margaritifera margaritifera. the foot is decreased in size through retrogressive development. France). and ventral surface. substratum in order to hold Mussels are buried in the quiet parts of the river bottom where it is covered with finer gravel and sand. about twice natural catfish sp. Lithogenes villosus in the rapids of The organs of rivers in Guiana.. 92. attaches itself to the substratum by means of threads from its byssus gland. disk-like thorax.g. attach several threads to the fast while they change their location. belong to different groups of Salientia (Fig. a second. is developed into a sucker.. Besides these. which are poor in lime because they ing flow over granitoid rocks. A mussel of streams in Africa and South FIG. British Islands. its shell it When the spring snail. Their threads are so tough that the larvae can float out into swift water without being carried away.3 m. and an adhesive. 3 Spines on the ventral surface are often helpful The tadpoles of tropical mountain with similar adaptations also brooks rivers and additions. an attachment by means of spun fibers 93). or Placostomus. as in many attached mussels in the ocean. The brook pearl mussel. 92). Byssanodonta. larvae of caddisflies attach themselves by means of a few threads. reattaches a thread to some fixed object and thus protected against being washed away. the umbo of . 9 17 ' 31 - also occasionally may be employed in running waters. and they are thus protected from be- washed downstream. The whole mouth of armored catfishes. and. of attachment are especially unique in fishes and tadpoles mountain streams.Mountain Brook Adaptations 377 mayfly nymphs have organs of attachment in the form of thickened and spiny rims on the tracheal gills. America. Finally. mouth. Gastromyzon. may be reconstructed into sucking apparatus. treats into is After Rauther. In thickened old specimens. There are only a few species in the fast-flowing waters of central Europe. and on fishes the fins also. these animals cling so tightly that they can be loosened only with great difficulty. Sucking mouth of the armored Placostomus from swift water. Germany. which are found in great variety in South America (e. BythineJla dunckeri. lower lip. Fig. size. This may occur in very different taxonomic groups. a few insect larvae spin threads for anchorage. which need a minimum current of 0. as is the ventral surface of the cyprinid. and larvae of black flies. in North Borneo. lives in brooks of north and middle Europe (Scandinavia.

Trichoptera.). c. western North America. except in progressive reduction of the fauna. etc. Perlidae. Original drawings by Fie. also widely distributed in much corroded by free carbonic is small ubiquitous Pisidium brooks. which emerge in the Modification of tadpoles tor lite in mountain streams. and do not contain de- veloping larvae again until fall. Bufo penangensis. Dodds 4 found that the species in higher altitudes . which sorts terrestrial animals and plants into different belts according to elevation. Staurois mantzorum. d. a f Hanu pijriens tadpole with unmodified mcmthparts. Or ton. spring in the lower stretches. The mountain ( The fauna of mountain brooks mainly consists of larvae of insects Ephemeridae. in his examination of the distribution of Entomostraca of the Colorado mountains. is less evident in running waters. The zonation of the highlands. Grace L. Malay Peninsula. west China. Even so. 93. have a reduced fauna in the warmer season. and in early summer in the higher regions. Such life histories explain the peculiar phenomenon that these brooks are most densely inhabited during the fall and winter months. Ascaphus truei. mouth parts enlarged to iorm an effective sucker in: b.378 these mussels is Communities often in Running Waters acid in the water.

either from the ocean up the rivers. That this fact is not simply dependent upon elevation above sea level is proved by the Aksai in the Tien Shan. besides the sparseness of food. meadows little plant material reaches the brooks as food. This has been proved to be true of the Cobitidae. streams. and the individuals are smaller. species of On- . Thus the amount of animal life decreases rapidly with an increase in elevation. and the algae are rather sparse at such a height. the number of treeless is reduced. while in larger rivers. above sea level on the southern slopes of the Tang-la. may be found in the circumstance that these stronger currents demand a much more strenuous use of the muscles of fishes in order that they may maintain position in the stream. as in central New York or in Illinois. in the Alps per- haps to 2800 m. Fishes ascend to varying heights in the mountains. In the Asiatic highlands two species of fishes have been taken in a spring with a temperature of 18-20 at a height of 4780 m. in the larger streams. thereby leaving a much smaller margin of food supply for growth. 27 An added reason. salmonoid fishes (Salmo salar in the North Atlantic. The average weight of all the trout caught in the Aar basin during 1913-1914. The average weight per unit is 30 greater. Migraor. numerous species of tory fishes going upstream include the sturgeons.Fish Migrations 379 tend to be found in higher latitudes in the lower-lying plains to the In the regions of perpetual snow and the adjacent areas of east.5 kg. however. more rarely. like the eel.. The trout.. which flows clear and quiet at a height of species 3000 m. compared with larger. above sea level and there contains just as large fishes as farther down. it becomes considerably larger. and for the tributaries alone only 132 gm. Fish migrations The migratory fishes form a unique part of the composition of the faunae of inland waters. These include fishes that travel during the spawning season. 18 The fact that the genus Nemachilus is represented by four species in these marginal regions of fish distribution perhaps depends upon the cir- cumstance that in these regions of very low barometric pressure the breathing of atmospheric oxygen supplements the oxygen supply. like the salmon. the numbers of fishes may be two as to three times as great per square meter of surface in the smaller. Salmo fario. for the Aar alone was 240 gm. reaches a length of only a few inches in the mountain brooks of Switzerland and a weight of only 0. In regions where the conditions in headwater streams are less severe. from the rivers into the ocean. and especially in mountain lakes.

000 fishes in one day/' The immense numbers of the whitefish Coregonus leucichthys and other species that come up the Obi and the Irtysch in the spring furnish opportunity for great draughts of fishes by the inhabitants of these regions.* The migration often occurs in great masses in rivers where civilization has not too greatly altered the environment. of all fresh-water fishes. spawn perish in the end from sheer and unexpected result of American fish tagging experiments. they are crowded against the banks. for many salmon remain permanently in the streams. such wheel could catch as many as 14. A. Chars (Salmo arcturus and S. besides these. Salmo coregonoides. and people. . whereby the them in their These studies have In the rivers of the far north such migratory fishes make up an important part of the fauna. have not yet been discovered. which flow to the Pacific from the Okhotsk highlands. and several species of shad. naresi) * Alosa alosa and A. bears. their backs pro! rude out of the water. and a trout. It is unlikely that any of these migrants ever return to the ocean. to penetrate farthest to the north. are sexually mature. instead of nets being used to catch One them. fishes are able to detect significant differences to guide 22 return migration. also shown that such fishes living in a long stream are physiologically different from those of the same species from a shorter river. while masses of others become the prey of birds. Only two species of salmonids in the rivers of northern Siberia are entirely independent of the ocean. 143 salmon were caught in one day in the year 1647. upstream migration of salmon 13 (Oncorhynchus) lasts from spring until autumn. even those that finally 13 exhaustion. some perish. the grayling Thymallus thymallus. the mass of fish going up is so enormous that.380 corhynchus Communities in Running Waters in the north Pacific. It is probably this migratory habit that enables the Salmonidae. finta in Europe. In the relatively short rivers of east Siberia. particularly on the Pacific coast. In the Rhine near Strasbourg. the keta ( Oncorhynchus kcta ) comes up in such great numbers to the brooks at their sources that the streams are not large enough to hold the fish. In the Columbia and Sacramento rivers of the Pacific coast of North America. is found in the evidence that salmon and fishes of similar habit have a strong tendency to return to the brook in which they were spawned when they in turn interesting An The mechanics of this process. large bucket-wheels are employed to scoop them out. sapidissima in North America. dogs. seven other species inhabit these waters. species of Coregonus in the Arctic).

Taylor & Francis: on aquatic insects: flies in 4. 1848. 1925. 135 figs. Hesse. 178. S." Univ. 8." Bull. Fischer: xii. W. BIBLIOGRAPHY 1." 11." in idem. Smith College: xii. 4:785-1094 (p. IV. D. Middendorff. Bauwstoffen voor eene Fauna van Nederland.. Mass. S. "The fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America north of the Isthmus of Panama. onder medewerking van onderscheidene geleerden en heoefenaars der Dierkunde. and F... such as the migration of redhorse and suckers in the Mississippi Valley. Richard. Jena. vol. M. 86 text figs. 3 figs." Pt. Corn- mentary textbook of fresh-water biology for American students. . P. Natl. The life of inland waters. 1872. 12. 9:1-168. 164 figs. 1889. Michigan. 1921. an eleIthaca. 603. 1899. "Ecological studies flies. and B. Needham. and caddis Dnfiem. Leiden. 3 pis. (p. C. Bur. 391 pis. 4 pis. stock: 1-438. Day. Flower. Hamburg. Brill: 3 vols. Dunn. Galtsoff.4:xiv. von. E. S. 1939. A." 1899:885-916. 1900. J. G. 13. Beitrage zur Kenntnis der Palingenia longicauda ( Ephemera Swammerdarniana) Elberfeld. list of the species recorded from those countries. Cornelius. 1853-1866. Dodds. Francis. 548. stone the Colorado Rockies. "Plankton entering the Huron River from Portage and Base Line Lakes. "Die Deutschen Siisswasserbryozoen." Ecology. "On a second collection of batrachians made in the Malay Peninsula and Siam from November 1896 to September 1898. J. 5 pis. 39:347-438. 82 34' N. 121). G. 1916.. 1924." Trans. Von den Antillen zum fernen Westen Reiseskizzen eines 6." Faune la Suisse. l:xviii. D. "Die Thierwelt Siberiens. S. T. 12 pis. the upper Mississippi. Idem. 5 pis. S. M. Evermann. Naturforschers. Zool Soc. 11:5-136. 5. Victor. Soc. 19 figs." Teil I.. 14. E. 1896. de 9. Altitudinul range and zonation of may flies. 613.mall streams. The migration of fishes from the seas to rivers* is paralleled in >ther situations by migration from lakes to rivers or from large to . (p. 47. and J.. l. Microscop. Fauna of British India. 87 figs. 16. 244 figs. "Fishes of Colorado. vi. pis. S. Abhandl.Bibliography 381 lave been caught even as far north as Grinnell Land. Ellis. 1843-1844. Ver. latiude. 1867. Franz. x. 162). "Histoire naturelle des reptiles et des batraciens. J. 1-4 :lx. T. Reise in den iiussersten Norden und Osten Siberiens. 59-60. Herklots. Colorado Studies. . Naturw. pts. A. 3313. Jena: iii. Museum. 1924. 2. 3. "Histoire naturelle des poissons. Hisaw. Fatio. 1882.. No.. Fisheries. 441. 6:380-390.. C... L. Am.0. London. 7 pis. 1926. M. Chandler. Northamp- 7. 15. K. Biischler: 1-38... 58:24-41. . F. Tiergeographic auf oekologischer Grundlage. 3: xii. Lloyd. ton. Fishes. R. 786. London. "Limnological observations in Bull. The salamanders of the family Plethodontidae. Kraepelin.. with a Proc. 956). 1887. 1913.. 17. Jordan.

33. S. Needhain. 184. F. 64:238-247. Noble. illus. Fischerei /. G. "Animal communities in temperate America. 1915. N. 280). Communities R. 1757. 1928-1929].382 16a. Schaeffer. Das fliegende Uferaas oder der Haft wegen derselben llten Augustmon. B. Surbeck. Limnology.. Ser. Berlm. 1734-1742. Y. 7. Imp. 22:551-570. (p. P. Ulmer. 31. Steinmann. Wright. 1907. R. Y. "Homing instinct in salmon. 9 14:408- 430. "Der Bergbach des Sauerlandrs. North Borneo. 1939. HydrobioL. and Am.. 46 figs. Paul. Biol. Exploration of Mount Kina Balu. maps (p. Fisheries Soc.. Microscop. 1914. 1902. J. Przevalski. T. lustrated in the viii. N. am Regensburg." Biol Zentr. 4. as ilChicago region: a study in animal ecology. A. 32. London. 21. figs.. Reisen in Tibet und am oberen Lauf des Gelben Flusses in den Jahren 1879 bis 1880 von N. Jena..." Int:rn Rev. Memoires pour servir a I'histoire des 267 pis. 27. The biology of the amphibia. "The Am. 317. 17). 260-266. 362. 30. illus. John. xiii. Unsere Wasserinsekten. 20. Naturalist. illus. Trans. P. Teil.. effect of stream conditions on lake plankton.. 1907. Heft 2:1-125. 281. 5 maps. Rev. 22. M." 238.. Soc. 26. 24. 1913. Thienemann.. Suppl. von Prschewalski. Gurney & Jackson: x. G. Georg. Meyer: 1-157. Am. "Quantitative of stream bottom foods. 22:228Geschlechtsverteilung bei Fischen)... Praktikum der Susstvasserbiologie. 5:xiii. E.. Rev.. 1884. N. royale: 7 vols. August. an der Donau. Reif. Whitehead... Paris. F. R. (p. 19. Welch." Intern. 1929. new ed. L. B. I. Shelf ord. FJuges. distribution." Trans.. 306 figs." Bull Grog. 174 18. Skorikow. C. 1908. "Die Tierwelt der Gebergebache. Biol Lacustre. A.. "Die Erforschung des Potamoplanktons in Russland. 112). 21 text figs. 1893. Gebriider Zunkel: 1-34.. 29. Woltereck. 41:351-354. "Plankton und Seenausfluss. 471. C. 1911. Soc. einr fiiunistischebiologische Studie. 2:30-163. Borntraeger: 23. McGraw-Hill: 577.. Hydrobiol. 1939." "A graphic method of correlating fish environment 1 fig.. 1931. [\ol. Scheer. 1:303-304. "Ueber die Ergebnisse der ersten Lfuchfischfapg-S+uiistik des Kantons Bern pro 1913/14 (Zugleich neuer Beitrag zur K'Tmtnis der Schweitz." Quart." 58:398-403. S. 32 pis. McGraw-Hill: xh. Costenoble: xiv. Paris. Leipzig." Ann. . 9 diagrams. Chicago. 1912. A. Quellc & 28. V. Reaumur. in Running Waters studies 1934.eiiunv. 25.. frontis. insectes. 17. und sonderlich auf der steinernen Brucke zu Regensburg ausserordenthch hailfigen Erscheinung u. K. Idem. H. de. Lechevalier. Biol.

An autochthonic river plankton can scarcely be said to exist.J. their increased content of dissolved substances and suspended detritus. The current in rivers often produces steep banks. the slope of the shores is more often gradual. the elements dissolved from the bottom tend to re- main. may be a segregated The favorable character of the environment for living forms de- creases wherever a river flows through a lake. and a wide border of plant growth supports a rich fauna. through the disintegration of dead plants the rest of the water is well supplied with detritus. The fact that the plankton of open water has more favorable living conditions in standing waters than in rivers also depends on the decrease of current. is limited. Standing water and almost self-sufficient habitat.O.e. i. The most favorable conditions are found in the slowly flowing lower course of the rivers. or have been brought in by tributaries are constantly being carried off to the sea. the river plankters are incessantly being carried to the 383 . it is greatest where the renewal is slowest. in lakes with outlets. In standing waters. the production of plankton is in direct ratio to the time required for renewal of the water. or have been set free by the disintegration of dead plants or animals. with their decreased velocity. Communities in Standing Inland Waters THE ABSENCE OF CURRENT IN STANDING WATERS ACTS IN SEVERAL WAYS TO determine the nature of the fauna. In standing waters. which are necessary for plant life and the quantity of which greatly influences the fruitfulness of the water. In a river the enrichment of the water with nutritive materials. the banks are scoured and their slopes increased by lateral erosion. on the other hand. Kofoid 3U states that.. All salts that have been dissolved from the ground. however. and those vital elements that have already been used in developing living forms also find their way back into the water at the death of these plants or animals. at least at high water.

Communities in Standing Inland Waters as nourishment where they perish and serve fishes. Aside only such animals as are buried in the substratum. this situation favors the volume than its in lake with 475 sql 43 the upper deep waters. because of their generally rapid current.' Only in very few do they exceed There is 1000 m. km. and the ratio of the depth to the surface area is certainly not fixed. and the amount of salts dissolved from the substratum. The shallow zone is is more extensive in relation to total area. and less in a pond than in a pool. the ratio of the amount of water to the extent 6f substratum greater in small. the transference of the spermatozoa to the eggs being assisted by the current. are every gradation of size in standing waters between Lake with 82. the smaller streams and brooks contain relatively fewer floating or free-swimming Smaller bodies of standing waters.384 sea. The depths vary similarly. 91 reports that the normal condition of Anodonta in standing water is hermaphroditic. of surface area. on the other hand. but from this depth there are all gradations to the shallowest puddles. and ponds. the fauna of the open water plays an important part along with that of the substratum. whereas those in flowing waters are bisexual. animals. especially of lighted bottom. if fairly permanent. for the surface area is larger here in relation to the the sun's rays. abundance of living forms. There is relatively less bottom area per cubic meter of water in a lake than in a pond. factor of extent its Aside from the nature of the substratum and richness in nutritive there are differences related to the ratio of water mass to bottom area. In standing waters. is also is much greater. and only rarely are they more than 400 m. or live in sheltered niches can hold their places in the current. The conditions of oxygen supply are more favorable in shallow waters.360 sq. and thus the of the water total vegetation is relatively greater. As was mentioned above. more favorable to life than the larger. other things being equal. and fertilizer. 80 In run- ning waters. or are attached to plants or stones. A much larger part penetrated by growth of plants. of surface area and its depth of more than . from Thus Weissensee Biological relations may vary decidedly in the two locations.. puddles. In Lake Constance km. An increase in the relative amount often means an increase in the of bottom area. The salts. the relationship is less favorable in deep lakes than in shallow ones. and Superior. pools. shallow waters.

km. which are richly fertilized. usually much less). able size and of a depth reaching 25 m.4 kg. and their place is taken by the warmer masses of stagnant water. and this process continues until the whole water mass has again acquired uniformity of temperature at 4.200 hectares (about 287. per hectare (respectively. 1. are reported from lakes in Canada. After long-extended cooling the whole mass of water reaches a uniform temperature of 4. per acre).7 Similar rekg. During the season when . the hand.2 and 2. and an much average depth much less than the maximum of 45 m.800 hectares (about 1.Temperature Relations 200 m.). per hectare (101 Ib. 113 kg. and 2. and sink. the density of fresh water increases with a decrease in temperature until it has reached its maximum at 4.000 acres). with 566. and amounts to only 1. consequently the surface layers become denser and sink until they reach a layer of equal temperature. The cooling of water occurs by radiation of heat from the surface. the water becomes stratified according to density. is much smaller. which is connected with the upper and has a surface area of 63 sq. when a covering of ice hinders any further giving cooling effect spreads but slowly to the deeper regions because of the poor thennal conductivity of water. 6 85 the smallest. sults Temperature relations is Temperature consequence of its of special importance for life in fresh waters. with an area ' of 31 hectares (77 acres) had the largest amount of bottom fauna and the greatest yield of fish. and Lake Malar. deep. and finally. such as Lake Vener. Among ponds in Holstein 5 all those rich in plankton are relatively small and shallow (not more than 7 m. when cold surface water with a temperature below 4 is the upper layers become denser as their temperature rises. Further warming then continues in the surface layers and penetrates the deeper water very the surface is warmer than 4 the slowly. of 116.4 Ib. Further cooling of the surface water causes it to expand again and become lighter. is 385 poorer in fauna than the lower lake.000 acres). the yield of the largest lakes.400. etc. and those of considerin plankton. which rise to the top only to sink again when they become cooled. As stated in Chapter 16. On the other off of heat. The relatively largest of amounts living forms are found in the small ponds (village ponds. and then with a further lowering of temperature the water expands again and becomes lighter. or more are invariably poor Of twenty Swedish lakes. warmed. this cold-water layer remains on top. In of high degree fluidity. per acre).

..e. They occur most regularly where the greatest changes of The convection Winter XII 1 Spring II Summer V! Autumn X Winter I Spring III Summer VI VII VIII III IV V VII V11I IX XI XII II IV V 70 Warming FIG. winter stagnation. i. temperate zones. . dotted when it is inverse. summei stagnation at fl-C and F-G. At a given time in autumn all at the surface. viz. This results in convection currents until a uniform temperature of 4 has again been reached. the water has a temperature of 4. 94). on the one hand. in the temperature occur. 94.386 Communities in Standing Inland Waters temperature decreases from the surface to the bottom. which continues up to the maximum of temperature. this is inverse stratification (Fig. Cooling Warming Schematic representation of the temperature condition? in a body of water in the temperate zone during the course of the year. Thermal conditions in standing waters are very strongly influenced by the change of seasons in the temperate zones. this is called When the surface is colder than 4. the temperature rises with increasing depth. Spring overturn at A-B and JE-F. and in the tropical or subtropical regions on the other. currents set up by the cooling and warming of of are water great importance in supplying the deep water with oxygen. direct stratification. if it continues to cool an inverse stratification sets in without creating con- vection currents. Then warming begins. autumn overturn at C-D. they are much re- duced in the polar regions. The isotherms are continuous lines when the stratification is direct.

The location of the thermocline or . tion of lakes on the basis of depth. below the level to which the daily convection currents extend. A discontinuity layer called the thermocline (metduction. Evidence that such less dramatic temperature changes tinuity layers is may indicate the presence of disconfurnished in these African lakes by accompanying in ionic content.Temperature Relations 387 The water is now in unstable equilibrium and wave action usually produces a complete mixing of the waters of the lake. a rapid decrease in temperature sets in. Consequently. of course. Thereupon the heating of the surface water goes on from the end of April or the beginning of May ( summer stagnation) until a renewed cooling again brings about equilibrium and the autumn overturn. 37 9 This establishes an important classificaparticularly in Lake Nyasa. exists only during the summer stagnation. Warming during the day in summer is followed by a cooling at night. however. The change from day to night also sets up convection currents. but twice a year it extends Between these two points the 4 to the surface. go down only to a below this the water is warmed only by means of conacts very slowly. certain depth. so that these upper strata differ but little in temperature. 9 8 Lake small and but Nyasa fairly marked changes in temperayika The thermocline is ture with depth are called thermoclines. two more subdivisions have been recognized in the hypolimnion. making what is called the spring overturn. which causes a sinking of the upper strata down to a stratum of like temperature. In contrast with the more definite epilimnion. it sinks deeper and deeper and at last defined by Birge's informal rule 14 as the depth zone in which the drop in temperature is 1 or more per meter of l)2 for example. 94). isotherm describes a curve that sinks lower in the in the summer than winter and above which the water is tification in summer and in inverse stratification in the arranged in direct strawinter (Fig. As a result of this an equalization of temperature among these strata occurs. in spring and fall. Such convection currents. with the beginis formed at this level. in Lake Tangandepth. which alimnion ) In temperate lakes the thermocline. changes in dissolved oxygen and Even when sharply developed. the thermocline tends to be more abrupt at its upper than at its lower limit. This rule often does not apply. ning of autumnal circulation disappears. fills Water at a temperature of 4 thus always the lower levels of the deeper basins.

and in shallow if lakes. alternately warm and cold (temperate . and 102. deep.Content 60 in 80 100 % of Saturation Temperature curves ( % of saturation. in less than 1. FIG. in Lake Tanganyika.5. they are exposed to mixing by means of the wind. The thermocline may be also differs in the same lake near the surface at a depth of 1 to 3 m. Madue Lake. the temperature is more equally distributed through their depths. 95. 10 35 or 8 considerably lower.) Temperature in C 8 10 12 14 16 18 20 22 24 2 4 6 40 20 40 0. particularly as regards wind action. Its depth in different lakes is indicated in Figs. has divided lakes into three types with reference to their temperature. often at 60 to 80 m. New of deeper lakes. and in Cold water is brought to the surface along many the shore of Lake Michigan by off-shore winds of a few days' duration during summer. 101. differs in different lakes in relation to latitude it as a result of changes in season and in weather. After Thienemann. is 1. especially. for Cayuga Lake. This type of temperature stratification applies to the open water In the shore regions of such waters.. ' (See Chapter 19. according to whether the surface water is always warm Forel 29 (over 4 in tropical lakes). 7 m. thus the greatest difference between the temperature of the surface water and of the deep water lakes is Lake Mansfeld. and Keller in Holstein. or more. 23 or somewhat deeper. ajid curves of oxygen cor tent ( ) in Lake York.388 Communities in Standing Inland Waters and nearby topography. ). 95. Ponirrania.

36 In running waters. even when the movement is slow. becomes so warm in summer is small. according to the regional climate. for their water sinks below the warmer water of the surface. or always cold (polar lakes). Lakes of the tropical type are widely distributed throughout the tropics and subtropics. deep (as.. which brings to the bottom water that has been oxygenated at the surface. Polar lakes always contain cold water with a temperature lower than 4 and have inverse thermal stratification except dur- become equalized. in that its type. on the other hand. but Lake Tanganyika and Lake Nyasa are 9 tropical in type as well as in location. without any movement of the water. The lakes at the southern base of the Alps. of overturn. even Lake Baikal (51 -55 N. Where there are no such tributaries. Stratification is always direct in tropical lakes. and in spring and fall equalizing circulations appear. so that it belongs to the temperate Deep lakes tend to be either polar or temperate.Oxygenation 389 lakes). the temperatures of the surface and the bottom and the stratification disappears. in summer they ing summer. 92 The convection distribution of currents are of considerable importance in the oxygen in standing waters. two periods . resemble tropical lakes. It is estimated that it would take 42 years for one molecule of oxygen to be transferred by means of diffusion. This is not true in standing waters. belong to this group.e. Conditions in temperate lakes are stated above. Lake Constance). in spring and in set the waters of temperate lakes in motion even to great depths. such as Lake Geneva. the characteristics are those of the polar type. i. though tributaries of lower temperature increase the oxygen content of deep waters in lakes of the tropical type such as Lake Como and Lake Geneva. depending on latitude. In contrast with these. the fall. when circulation occurs. in the winter polar lakes. the shallow Lake Enare in Lapland. north of the Arctic Circle. for instance. Polar lakes occur in arctic regions and in high mountains near glaciers. latitude) and Lake Telesky in the Altai belong to this group. but wherever the ratio of the surface area to the volume of water deep lakes in subpolar regions. are usually of the temperate 1()0 or tropical type. from the surface to the bottom of a lake 250 m. and really shallow lakes are not deep enough to become thermally Oxygenation stratified. constant mixing takes place. waters are directly stratified. Relatively shallow lakes.

Our general discussion of the fauna in standing waters will be based primarily on that of lakes. there is no lack of is oxygen at the bottom.demarcations between these groups or their corresponding waters. no waters fall sharp .. shallower. Communities dependent on the bottom are said benthos in oceans and to the pedon (pedonic) in belong to the lakes. Lake Cayuga). in which the shore vegetation does not extend to the greatest depth. A lake may be defined as a continuously closed. Lake of normal saturation content less. In waters whose substratum is mainly composed of mineral elements the oxygen content near the bottom does not sink below 70% of saturation (Fig. which is rich in disintegrated organic material. contain open water and have a limnetic region. of course. but the limnetic . especially in shallow depths. Wherever the bottom is covered with ooze. 95. more evanescent ponds and pools on the other. Lakes. The region would be that of the paralimnion. In this respect conditions in a lake' are more like those in the ocean. often only 40$. the is oxygen content. 95 ). or. paralimnic. stagnating mass of water found in a depression of the earth. If need is felt for further sharp- ening of this terminology. none at all (Fig. in deepest layers just above the floor. in contrast with ponds. much reduced Madue). or (Fig. and a new term. but the oxygen content of the deeper water is depleted because of consumption of oxygen in that region. 95. 95. There are. 83 In such waters the bottom fauna which wave In shallow greatly influenced. and in which. may be used for the corresponding community in lakes. so that in the The decrease lakes. is left. action extends throughout.Animal communities of standing naturally into two groups those of the deeper and more permanent lakes. therefore. Animal communities in lakes Communities of organisms or shore are called pelagic free from direct in the dependence on bottom if if ocean and limnetic to in lakes. never directly connected with the ocean (Forel). on the one hand. The com- munities related to the shores are said to belong to the littoral community in both oceans and in lakes. there is a central area free from rooted plant growth.390 Communities in Standing Inland Waters During the summer the surface layers are well mixed and well supplied with oxygen. of oxygen content begins at the thermocline (Fig. and of the smaller. Lake Keller). littoral may be retained for such communities along the shores of the ocean. therefore.

the paralimnic zone for instance. floor. Where the slope is gradual. and reeds and growths of rushes and sedges. 391 much The shore zone The pedonic region far as the green plant is composed growth on the lake zone beneath the limnetic region. 96. or in the Scottish lochs. are into several parts. stemmed grasses high of swamp plants. After Brutschy. and this is the greater when the amount of plankton present in the lake is smaller. (Fig. found the thickly growing emergents. where the slope is steep. the . This depends primarily upon ^r Nymphaea Schoenoragmites * o Chara Najas plectus Potamogeton Cera to phyllum Myrio- phyllum FIG. sometimes but usually not exceeding 6 to 12 m. as. quantities. contains by far the richest and most diverse fauna It furnishes substratum for animals on its floor and on its plant forms. The paralimnic zone may be divided in the shallowest waters. afford nourishment. certain species The gentler the slope of the shore. such as strip pondweed (Potamogeton) and water lilies (Nymphaea). Elodea. tion extends varies in different lakes. The shore region in lakes.. in it many places in is wide. Finally there is of Isoetes. down to about 30 m.The Shore Zone region in general does not exceed the pedonic region nearly so as the pelagial exceeds the benthal in the ocean. etc. Schematic diagram of one type of plant growth at the edge of a lake. Chara. a zone of submerged plants. 96). the transparency of the lake water. Lake Lucerne. which extends as and the deep pedonic The depth to which shore vegetaof a shore zone. and liberate oxygen in great First of all. the plants supply shelter from enemies and protection against wave action. may be entirely absent. then follows a a mass turn in which grows among more of strictly aquatic plants with floating leaves.

We have preserved the familiar Corethra for the purposes of this book. whereas in the open water near the surface the temperature is only 2. For this reason ice first forms along the shores in winter.* Libel- lulidae) congregate and undergo metamorphosis in such warmer regions. on plantless shores where strong wa"es beat when the wind is high. there all the loose particles on the bottom are washed away and a surf shore line of gravelly or sandy beach develops. No plants can grow where waves driven by prevailing winds break strongly against the shore. Fishes congregate where the bottom food supply is plentiful. for the cooled surface waters cannot set up a circulation like that of open water. in Lake Constance a yearly average variation of 2. Many insect larvae (Corcthra. This is especially true in mountain lakes where the periodic increase and decrease in size of tributaries fall from glaciers or melting snow cause a marked rise and of water level. A harmful factor for the fauna of a littoral region is the regular variation in the water line in many lakes. This is tus feeders and furthers especially advantageous to the numerous detritheir increase. Even in the Great Lakes there are exof tensive irregular variations in lake level both from temporary seiches and over periods of years. where a temperature of 17. Cooling of the water likewise occurs more rapidly in the shallow shore waters. Because of the closeness of the lake bottom. The temperature factor is subject to greater change in the paralimnic zone than in other regions of lakes. Only that part of the shore fauna which can * resist or evade the = Chaoborus. rocks and pebbles are tossed about.2 may exist under the noonday sun. providing thus for the development of detritus and the occasional stirring up of these nutrialso grinds tive materials. with its own peculiar sparse animal life. .2 m. with a maximum in the lakes of more than 3 m. is fatal to tected from wind.5 98 . icefloes are dashed against the banks. and experience in fishing shows that exposed shores may furnish a richer catch of fishes than shores pro73 A region of strong breakers.. The breaking of waves against the shore not only kills organisms but up their excrement and remains. This is especially noticeable on shores with southern exposure. und in wintei. most forms of life. the water warms much more quickly than in the region of open water.392 Communities in Standing Inland Waters wider these zones are found to be. A similar condition occurs the plains. hovsever.

a. or that stand drying that (threadworms. animals desiccation by encysting. tardigrades. in winter the oxygen liberated by photosynthesis collects in large bubbles under the ice and can be utilized by Contrary to a condition frequently found in air-breathing insects. Sessile FIG. or. : X or slow-moving animals. submerged tree trunks. insects. solid objects such as reeds. and water mites. digging in the mud with its rostrum. the aquatic insects are seldom limited to definite plants. partly in the leaves of plants. forms this habitat the center Above and mussels. finally. and rotifers are abundant. mud (Fig. In the region of profuse plant growth. 25. annelids. Littoral mud-inhabiting cladocerans a. as well as ladders used in climbing to the surface of the water for air. partly among the There are along with isopods and amphipods. such as crustaceans. leeches. after Fri Vavra. 97. snails numerous all. b. lliocnjptus sordidus. being especially plentiful where the growth of Myriophyllum is thickest. Fresh-water sponges and occasionally many kinds of Bryozoa grow on stones. are absent. resting places. many rotifers and copepods). and . which retreat or advance with the water line. 98 which find nourishment and of life for both adult and larval insects. 97).The Shore Zone 393 unfavorable conditions can live in such parts of the paralimnic zone as are dry a part of the year: animals with powers of locomotion. b. terrestrial insects. animals of amphibian habits. such as frogs and many snails. Protozoans. flatworms. and mussels. bryozoans. Rhynchotalona falcata. such as sponges. 45 With them live a host of Entomostraca. after Herr. on the bottom. with the animals that prey upon them. in and among the leaves and is a diverse and important group of plant and detritus feeders. like many protect themselves from Protozoa.

: the Chiemwide whose shallow and Bavaria). 98)*' to Numerous flatworms hide under Ibs. covered with plants. Euspongilla lacustris from quiet water and from mo ing water. without a paralimnion. 1 .394 Communities large in Standing Inland Waters A number growth. high (Fig. . of fish 71 per hectare. from rather large variations see (in whose shallower water suffers produces 8 kg. the fertility of in water level. attached After Weltner. The Konigssee (in young Bavaria). 5. of fish per FIG. neighboring streams. produces annually 1 kg. paralimnic region is well Lake Constance.. 3 It is unsafe. * 0. where no plants are found. however. The animal life of wave-beaten rocky shores. Fresh-water sponges grow on the stones in flat. to a floating thread. to generalize too confidently at this point.- ^S^Hv . for this reason Thus it hapfishes are plentifully represented in this region.9. slab-like crusts.* These relations prove to be similar in the United States.* i hectare. has a faunal composition much like that of swift water in 96 A close observer always finds teeming life. of of that the paralimnic plant growth may development degree pens have a marked effect on the fish fauna of a lake. produces about 20-25 kg. per acre. 98. . and of fishes find nourishment in the region of plant especially a favorable spawning place. while in quiet bays they develop into much branched stems often up to 30 cm.3. the stones and 18 22 respectively.

into which no light rays penetrate. normal to the arrow. are small and and convex. also occa- sionally the shield-shaped snail. thin-shelled. truly abyssal stratum. Steep shores are poor in life. Nyasa. ftuviatilis (Fig.The Deep Pedon 395 (Dendrocoelum. long. which in turn produces a reaction on the softer parts of the body (Fig. L. The mussels Unio of the plantless Anodonta (Fig. The amount of light that penetrates into the deeper waters is much reduced. bodamica from moving water. and in the brackish Caspian Sea. Mayfly nymphs are also much depressed. sucker-like foot. There are associated modifications of color in both . comparable with that of the sea. beautifully colored forms found in the quiet waters Lake Constance. is widely distributed. those in the Waldsee of bays. Herpohdella) attach themselves By means of their suckers. with relatively thick shells often considerably corroded at the umbo. Ancylus shores. stagnalis from quiet water. 49 to The snails of the genus Limnaca change form in the moving waters of the lake 19. b. The deeper regions of lakes have other peculiarities besides the lack of plant growth that contribute to the unique compositions of their fauna. is lacking in most inland waters. In lakes. I imnaea stagnalis var. for instance. with its wide. in contrast with the large. flatly depressed leeches (Glossiphonia. shore because of the constant pull to which their shells are ex- posed. stagnalis from a pond. of the lake bottom it is makes up the deep pedonic re- In biotic characteristics communities of the paralimnion. L. the snail Goniobasis. 99. the growth form produced by a strong growth of filamentous algae on the shell. beyond which the shell is weighted upper Wiirttemberg. a. Tanganyika. of in The Anodonta FIG. Planaria. and Posso. up cm. become 9 cm. 89). which hinders its motion. The deep pedon The remainder gion. producing growth similar to a (after Voigt). c. 99). Polycells). 70) alike. extensive development of such a stratum is found only in Lakes Baikal.6 with cement. ders are occasionally characterized not sharply marked off from the Those parts lying nearest its bor- A by an abundant molluscan fauna. many caddisfly larvae weigh down their cases with heavier stones.

carry many small plankton animals with them into the deeps and bury them. and no variation of sub- stratum. No influence of increased water pressure on the animals living on or near the bottoms of lakes has so far been demonstrated. Planaria alpina). no change of temperature. this is inhabited by an exceedingly sparse fauna. Because of this the floor is evenly covered with fine mud varying according to is not less than 4. cyanophyceous mud. the variation 100-200 m. The dead bodies of plants arid animals living in the open water. black vile-smelling muck rich in hydrogen sulphide is produced. a sterile. the excrements of fishes. scarcely any change in the no movement of the water. is low forms at depths of 1(U 1. and Such sediments.396 Communities in Standing Inland Waters limnetic (see below) and bottom forms. which is usually organic. the rest When more these organic elements are disinfected by the prevalence of humic acid and are thus kept from disintegrating. about Stenothermal animals. and bloodworms (Chironomidae). Where rapidly flowing streams of mountains or foothills contribute masses of mineral sediment. Temperavery constant in the deeper pedon as compared with that near the shore. although the evidence is not yet fully collected from lakes Baikal and Tanganyika. otherwise black (Polycelis nigra. more or less solid. These continually engulf the mud and give off small mudballs from which the largest part of the organic material has been exis finally disintegrated by the action of bacteria. organic material sinks to the bottom than can be consumed by the deep-water fauna and the bacteria. and the products of disintegration of shore plants sink to the bottom and form a layer of detritus. For this reason many periodicities common in the biota of shallow waters are not evident. and in especially pronounced deposits one can identify different sorts. are represented by pale yelture in the depths of the lakes in the Alpine foothills. such as diatomaceous mud. its origin.7 is and not more than 10. adapted to only the cold. lakes sometimes excessive in amount. and chitinous mud. accordingly the mud of the lake bottom varies with the various compositions of the plankton. with a few eurythermal forms. which becomes the food of innumerable pea clams (Pisidium). slimeworms (Tubificidae). Bottom mud in all deep comes primarily from decaying plankton. terrigenous mud becomes deposited on the bottom. There is no marked movement of water at greater depths. find this a favorable habitat. there is amount of light. Peat develops where tracted. The planarians. a resting period during the winter . In Lake Lucerne the temperature at depths of 30 to 200 m. Great uniformity of environment prevails at the bottom of deep lakes. finely grained.

partly also of species limited to the deeper parts of the lakes. in accordance with the conditions of temperature. A common characteristic of all these deepwater rhizopods is that they are larger than their relatives in shallow water.. the larvae of biting gnats. 5750 have been recorded from 10 liters of few Bryozoa also grow in deep water. widely distributed. The largest part of the fauna is composed of rhizo- pods. Where triclad flatworms occur in the deep regions they are reduced in size.Animal Life of Deep Water 397 does not occur. Slimeworms (Tubifex and may be the prevailing forms. 104 Animal life of deep water The deep fauna of lakes consists of ubiquitous eurythermal forms and stenothermal cold-water forms. e. as is Planaria //. pods. 110 /. Plagiostomum lemani (as in Lake Constance and other Alpine border lakes.g. and in Lake Madue in Pomerania down to 40 m. This is especially noticeable in Cyphoderia ampulla (measurin deep water. Tubificidae. especially in the deep lakes.. Of the flatworms a number of Rhabdocoela are generally characteristic of the deeper parts of lakes. Niphargus puteanus. e. often in surprisingly large numbers of individuals. Pisidium.. in shallow water). the pea clam. 307). or in summit lakes of the higher mountains). ostraFredericella distributed widely a are cods.g. The slimeworms place in numbers in the bottom fauna. and the cave amphipod. there are species of Canthocamptus especially. the sultana. (Tubificidae) take first . and several speA number of interesting cold-water isopods and cies of Cyclops. mud A amphipods live in deep water in addition to ubiquitous species in the border lakes of the Alps. Crustacea are common. a condition similar ing 200 that in to some deep-sea forms (see p. which is and relatives) are common individuals of various species its adapted to low temperature. large number of genera..g. Rhizopods occur in a rather large number of species. shows no clearly marked rings of annual growth but has a uniformly devel- oped shell. and the small bivalve Pisidium. is important. Corethra. in the deeper regions. alpina. As many as 8000 individuals of these forms may be found on a single square meter. of the coperepresented by from 32 m. The ubiquitous Dendrocoelum lacteum is an example. with chironomid midges next. these consist partly of ubiquitous forms. the occurrence of blind forms such as Asellus cavaticus. especially in the deep pedon of Alpine border lakes. The fresh-water polyp Hydra and the sponge Spongilla are found in rather deep water. chironomid larvae. in the Teufelsee (in the Bohemian Forest) down to 25 m. e.

palustris. i. gill- among which Bythinia tentaculata goes down to 12 breathing 72 In the Alpine borin. however. and only facultative anaerobes such as some protozoans and chironomid larvae can survive as permanent residents. forelia. This produces a severe selection. mussels and gill-breathing In Lake Ratzeburg. 104 They do not rise to the surface to paralimnic relatives do.. profunda. palustris are eurythermal. especially because of a rise in . and exhibits a great variability. in contrast with 7 snails.e. The fauna of the deeper pedon evidently originated nion and probably is constantly restocked from Most of the abyssal animal species also occur in shallower water. some pulmonates of the genus to life in the depths. The tiny bivalve Pisidium. in mountain lakes they are also found in the shore zone. for instance. as for instance in breathe as their Limnaea have become adapted Lake Geneva. In lakes with a thermocline the hypolimnion typically becomes oxygen-deficient and even oxygen-free. and Valvata piscinalis var. In Lake Geneva three forms Limnaea occur: L. at least in limited regions during summer stagnation... is very sparse. stagnalis and L. L. which has lusks of the brought about a separation into a large number of species. that are found in the deeper regions but are lacking in shallow waters of the same lakes are not specifically abyssal animals. It may be supposed that they formerly similarly inhabited the shores of lowland lakes. Thus the fauna of the deeper waters of Scottish lochs.. ovata and L. antiqua to 18 m. Those species.398 Communities in Standing Inland Waters upon the atmo- Most mollusks of deeper waters are not dependent sphere for their supply of oxygen. and L. and Bythinia occurs as deep as 60 m. for their descendants raised in an aquarium revert to the original type and again make periodic of such It excursions to the surface for breathing. but fill their lung space with water from which they take the necessary oxygen. on the other hand. and only because of a change in environmental conditions. even the plentiful pisidia. like that of their paralimnion. ovata and Planorbis albus go deeper than 3 m. abyssicola is a variety of L. has been proved that the first two are varieties of L. For this reason the amount and the composition of the deeper pedonic fauna depend on the type of paralimnic fauna of the lake in question. der lakes Valvata goes down even to 64 m. In a number of Alpine border lakes. lives in depths greater than 200 m. abyssicola. in the paralimthe same source. and L. whose water contains a large supply of oxygen in the deeper parts. ovata. L. of pulmonates only Limnaea snails. auricularia do not exist. All moldeep pedon of lakes are stunted. deep-water forms of the stenothermal warm water species L.

The conditions of the lake abysses.). species. 11. They Lake Lunz 160 individuals of the nannoplankton were counted to every 3 of those caught in nets. for it region that distinguishes 650040002100 . the dwarf or nannoplankton. especially in the spring. The plankton of inland waters consists of plant and animal forms. also eat diatoms . there is a of the the communities waters pond plankton. in Amounts of rotifers Saxony. inland lakes such as Lake Baikal and Lake Tanganyika (see Chapter tive of new 6 and p. ) in waters ) and of nannoplankton ( After Dieffenbach. forms that require organic food material. 29. . 850 250 SOIL 18..The Limnetic Fauna 399 temperature. 100. are partly photosynthetic and partly. 31. ( 10. 24. 410 ff. Large plankton animals. 4. some of the plankton animals are so tiny that they escape being caught with ordinary gauze nets and can be separated only by filtering or centrifuging the water. very deep. 5. though open of bodies of fresh water find their typical development in lakes. and in spite of the immense difference in size. have they been crowded into the depths. VI. 2672. X. Some of these organisms. Temperature curve ( which small areas of open water without vegetamay exist occasionally. 19. though to a less extent. 17. 63 Many plankters are restricted partly or entirely to the nannoplankton for their food. N /--A ." YD. The is limnetic fauna of The fauna open water is just this constant existence of a limnetic especially characteristic of lakes. YDJ.. from June to October. Fie. 7. but in which there is no continuous widespread region without plant growth. As in the ocean. 21. ). 25. DC. the former furnishing the foundation for the existence of the latter. the mass of which sometimes controls their numbers is (see Fig. have seldom proved produclived. 14. 100). the yearly average volume of the are so plentiful that in nannoplankton three times as great as that of the larger plankton. of course. consequently. since even the deeper regions are relatively shortSpecial abyssal forms exist only in the very old. 14. tion Alperceptible transitions make a sharp demarcation impossible. Imin them from ponds.

in contrast with small fertile village 77 02 On the ponds that may. however. region. the . m. small lakes richer than large ones. of ' amount of plankton in the Dobersdorf pond (Hoivaries between 136 and 3977 cc. only mals and plants in the plankton. whereas Bosmina longirostris is is permanently so only in the smaller bodies of water and The same is true of a number The amount and composition 10 only casually limnetic in true lakes.. only 11 occurred 10. of other Cladocera and of a fc rotifers. exceptionally. a fact explainable by the peculiar adaptations demanded by a purely limnetic life. over are or that receive other plentiful nutrient materials in sterile mountain richer plankton than those of a relatively granitic r> limestone. and in stein) Lake Plon between 13 and 424 cc. and a few rotifers. by the amount of dis- solved nutritive material in the water. above all. and of Moreover. limited. as well as in the same body of water at various depths and at various Very poor ponds or deep. p. 213). of 67 crustaceans known from Lake rotifers. in. a number of 95 Besides Crustacea and rotifers. cc. netic. during the course of one year.400 Communities in Standing Inland Waters and other of the larger in the plankton is algae. This is one reason why shallow waters in general are richer in plankton than deep waters. this is indirectly also responsible for the total amount of plankton present in a body of water (cf. etc. is truly limnetic. Corethra plumicornis (Fig. To the truly limnetic animals njay be added a numler of more or Bosmina core go ai is limless irregular visitors in the open water. cold lakes often contain only 5-10 animal plankton in 1 cu. of ani- mals only the species of Diaptomus among copcpods. Of a total of approximately 150 35 Balaton. As the development of plant elements among other factors. only a few species are sufficiently prevalent to determine its character. only 8 were limnetic. water mites furnish a number of species of limnetic forms. Of 66 phyllopocls of southeastern Germany. cladocerans. in 1 cu. Animals of open water are considerably in the minority compared with their relatives near the shore. 104). Only one insect. as limnetic forms. contain 1500-2500 cc. of water. times. w of the plankton are very variable according to the prevalent environmental conditions in various locations. among the numerous forms of ani- in species of plankton organisms of the Danish lakes only a few appear such numbers as to create a monotonous animal plankton. the mean other hand. why those lakes that are situated over fertile ground.. the so-called phantom larvy. The annual yield of plankton in the well-studied Lake Mendota (Wisconsin) is of the order of 5 tons of dry matter per acre.

Two different types of phytoplankton can be distinguished in temperate regions: in one of them diatoms are most common." is 401 15 Freshapproximately 200 Ib.. but in summer it is a bluish-green. about 12.The Limnetic Fauna standing crop zones. Perrennial plankton forms. most copepods. most prevalent in the cold northern or Alpine lakes where blue-green algae. in the lakes of Mendota the March minimum is almost one-fourth in winter.. many Bosmina. Thus the ratio of winter to summer plankton in north Germany 1:20. in fact. 1 1 ''" of various plant and animal species occurs at different times. are dis- tinguished from periodic forms that occur only at definite seasons Those without rest periods belong to the perennial of the year. is a sign of scarcity of plankton. e. in the lakes of Switzerland and the French Jura. species of which occur in unusual numbers. Examples of periodic . group. Many animal forms. The color of the water in colder lakes. are almost totally absent. 103 or in the different Alpine border lakes. and rotifers such as Anuraca cochlearis and Asplanchna priodonta. with few exceptions. the more turbid and discolored the water The tint depends on the composition of the plankton." In Lake of the April maxidefinite in the compothe course the of changes year During sition of plankton arise because of the fact that the breeding season mum. e. The optimum for most fresh-water diatoms is at relatively low temperature.g. is yellowish-green (because of diatoms and Ceratium). therefore. water plankton r