Field Tests of Theories Concerning Distributional Control

Author(s): Joseph Grinnell

Source: The American Naturalist, Vol. 51, No. 602 (Feb., 1917), pp. 115-128
Published by: The University of Chicago Press for The American Society of Naturalists
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No.602] SHORTER ARTICLES AND DISCUSSIONS 115

FIELD TESTS OF THEORIES CONCERNING DISTRIBU-

TIONAL CONTROL'
THE conditionsof animal distributionand the causes of these
conditionsare facts which concern intimatelythe problemsof
the persistence and of the evolution of species. The present
writerbelieves that the field naturalist is in a position to con-
tributein,large measure toward the solution of these problems,
and it is the purpose of this paper to show how comparative
studies in the distributionof species may throwlight not only
upon the nature of the environmentalcomplex,but also on the
relativeimportanceof its various componentfactors.
Some simple facts of distributionwhich are of commonobser-
vation, and which were early recordedby the systematiczoolo-
gist, are: (1) that each animal occupies a definitearea, that i1s,
has a habitator range,whichis distinctiveenoughto be included
among the charactersof the species and describedalong with its
habits and the features of its bodily structure; (2) that some
species (and even some of the highersystematicgroups) range
widely,and cover great extentsof country,while othersare ex-
tremelylocal or restrictedin their distribution;and (3) that,
notwithstandingconsiderablevariation in this degree of distri-
butional restriction,many species (or highergroups) are found
nearly or entirelyto coincidein range, so that sets of species,of
varying ranks, may be recognizeddistributionally,as constitut-
ing realms,zones, faunas, subfaunas,associations,etc.
Perhaps the most prominentdelimitingfactor,and the one
which has been emphasizedthroughrepetitionin the early sys-
tematicwritings,is the obvious one of physical barriers-repret
sented by bodies of water in the case of the terrestrialspecies
and by land in that of the aquatic. The majority of animals
inhabitingislands and seas are specialized in such a manner as
to be hemmedin by the limits of their respectivehabitats. In-
dividuals oversteppingthe barrier in either case are subject to
promptdestruction. This obvious type of distributionalcontrol
has always been and will remain an importantone for consid-
eration; but with the acquisition of detailed knowledgeregard-
ing the distributionof animals on large continental areas,
naturalistshave been led to propose many other factorswhich
l Contributionfrom the Museum of Vertebrate Zoology of the University
of California.

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116 THE AMERICAN NATURALIST [VOL.LI

have seemed to them to prevent the random and unrestricted

spreading of animals over the surface of the land. The fol-
lowing is a list of the factorswhich various writershave nomi-
nated as affectingthe distributionof the highervertebrateani-
mals. This list is completeonlyto the extentthatmy own exam-
ination.of the literature is so. Many of the items have been
found in dissertationsupon bird migration,which is, of course,
but one phase of the general subject of distribution.
Vegetation.
Food supply,kind and quantity.
Rainfall.
Humidityof the air (relative or absolute).
Wetness or drynessof the soil.
Barometric pressure, or altitude.
Atmosphericdensity.
Safety of breedingplaces.
Availability of temporaryrefuges.
Water (to land species).
Land (to aquatic species).
Nature and availabilityof cover,or shelterfromenemies.
Nature of the ground (coarse or finesoil, or rock).
Insolation, or light intensity.
Cloudiness.
Temperature: in general; mean annual; of winter; of period
of reproduction;of hottestpart of year.
Interspecificpressure,or competition,or race antagonism.
Parasitism.
Individual, or racial, preferences.
It is at once plain that some of the items enumeratedare ex-
tremely complex, and that the most superficial analysis will
show some duplicationamong them. For example,the factorof
vegetationas influencingthe distributionof differentmammals
resolves itself principallyinto the elementsof food-supplyand
shelter,and, subordinatelyin most cases, into those of tempera-
ture,humidity,and nature of the soil. As someof the suggested
factors may really never functionin any vital degree as sup-
posed, the total number of really critical factors is probably
smaller than the total of the items just listed. Time could not
here be taken to discuss the intrinsicnature of each elemental
factor,even if the writerwere equipped to handle such a variety

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No. 602] SHORTER ARTICLES AND DISCUSSIONS 117

of subjects; for such a discussionwould in most instanceslead

directly into physics and chemistry,and into a study of the
physiologicalprocessesof the animals affectedby each of these
factors. I should,however,like to dispose at once of one of the
"factors" listed, and which I hear and see repeatedly cited as
a cause of restrictionin distribution-particularlyin that of
birds.
Many people claim to see in the facts of distributiononly the
operationof a preferenceon the part of each animal-by virtue
of which,if a heterogeneouslot of animals were introducedinto
an area presentingdiverse conditions,each species would choose
its "natural" surroundingsand rapidly allocate itselfin a nor-
mal way. I grant that such a choice would almost certainlybe
made. In fact the hypothesisis being proved continuallyall
over the country in connectionwith the migration of birds.
Scores of species travel north in the spring to countriesfor a
preceding interval unoccupied; and while, roughly speaking,
they travel together,and arrive together,they segregatethem-
selves,immediatelyon theirarrival, and repair to separate sorts
of ground, each species by itself: the pipits to the prairie, the
water-thrushesto the streaniside thicket,the black-poll war-
blers to the spruce forest,and so on. We have here an obvious
choice exercised in the selection of habitats. But does this
segregationof species by exerciseof "individual preference" in
a uniformdirection change the nature of the problem in any
fundamentalway? Should we not here recognizemerelya char-
acter in the cerebral equipmentof each race, which,like every
externalpeculiarityin its structure,is in considerablemeasure
the result of protractedimpress upon the organism from the
environmentalcomplex of factors to which the race has been
subject throughpast time? There is no otheradditional factor
than those environmentalones (plus the intrinsicfixednessof
the species, within certain limits of plasticity,and the "evolu-
tionarymomentum") to be called into account.
As to the mechanismof geographiclimitation,the adjustments
to the various critical factorsare inevitablyforeverin process,
thoughreduced to a minimumat times of slow environmental
change. The refined method of individual "preference" or
"choice" is superior to the wasteful process of wholesale de-
structionwhich would be experienced by individuals finding
themselvesout of place as the result of a haphazard selectionof

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118 THE AMERICAN NATURALIST [VOL.LI

locality. The frontierindividuals,those on the margin of the

habitat of the species, may not prosper as greatly,or reproduce
as thosein the metropolisof theirspecies; butthey
as prolifically,
certainlydo not, as a rule, beat themselvesto death individually
against their limitingbarrier,of whatevernature it may be.
To resumethe main topic of this discussion,I shall attemptto
show that it is possible fromfieldobservationto indicate in the
case of certain species, some,at least, of the factorswhich con-
trol their distribution;and furtherthat we who live in Califor-
nia have splendid opportunitiesto gather and examine data by
means of which the general laws of animal distributioncan be
determined. An area within comparativelyeasy reach presents
a wide diversityin topographicand climaticfeatures. Occupy-
ing this area is an abundant complementof the highervertebrate
,classes. Withinthe politicallimitsof the state,systematists now
recognizethe presenceof 388 species of mammals,543 of birds,
79 of reptiles and 37 of amphibians. We have plenty of ma-
terial to work with. I shall proceed to discuss a few selected
species -aboutwhich we seem to have knowledgeenough to war-
rant provisionalinferences.

THE CASE OF THE OREGON JAY

The Oregon jay (Perisoreus obscures), a close relative of the

Canada jay, or whisky-jack,occurs in California only in the
northernthirdof the state. Even thereit is verylocal in its oc-
currenceand absolutelynon-migratory.On the Warner Moun-
tains, Modoc County,it ranges fromthe highestparts down to
7,000 feet altitude. On Mount Shasta it ranges fromnear tin-
berline down to about 6,000 feet altitude. It is absent for a
long distance to the west, throughthe Trinity mountainmass,
but it recursalongtheseacoastof HumboldtCounty,withinfifteen
miles of the ocean. And here is the curious point: along this
coast strip it does not range higherthan 300 or 400 feet above
sea level, althoughthereare mountainsnot far inland whichrise
to an altitude of several thousand feet. Let us look into this
case for the purpose of determiningthe factorsresponsiblefor
this interruptedrange.
The Oregonjay, like mostmembersof the crow family,is not
restrictedin diet. It eats a great varietyof both vegetableand
animal substances; its food varies in character according to
season and local conditions. The supply of any particularkind

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No. 602] SHORTER ARTICLES AND DISCUSSIONS 119

of food is not likely,therefoore, to be a controllingfactorin its

distribution.
The bird is a forestdweller. Its equipmentas regards man-
ner of flightand course to take in case of attack by enemiesis
adjusted to a foresthabitat, and nowhere within the writer's
knowledgedoes this jay extend its range beyond the limits of
woods of some sort. Although somewhatpredaceous itself, it
has regular enemiesamong hawks and owls, for protectionfrom
which it makes use of forestvegetation. This factor of forest
cover,then,mustbe countedas essential. But the range of the
bird is not continuouswhereverforestsextend.
In the interiorof California it does not descend below a cer-
tain altitude. Now three other factors in its distributionare
quite obviously connectedwith that of altitude, namely, baro-
metric pressure, atmosphericdensity, and temperature. But
when we take into account the fact that the Oregon jay exists
at or close to sea level around Humboldt Bay, the firsttwo fac-
tors,thoseof pressure,and air density,are instantlyeliminated,
because of the obvious fact that the bird successfullymaintains
itselfin localitiesof widelydifferingaltitudewherethese factors
are thus extremelydiverse.
With referenceto temperature,we know withoutrecourseto
instrumentation that there is a decrease upwards at an average
rate of 3 to 4 degrees F. per thousand feet. If, then, the bird
is limited downwards at a critical point, the inferenceappar-
ently follows that temperatureis the determiningfactor, and
this conclusionis inevitable if we consider only Mount Shasta
and the Warner Mountains. But the bird's occurrenceat Hum-
boldt Bay complicatesthe problem. In orderto reconcilethese
facts of distributionwe must look into the situationwith refer-
ence to season. On doing so we discover that the home of the
Shasta and Warner jays is subject to severe winterswith heavy
snow, very much colder than the winters at Humboldt Bay,
where the climate is equable and snow rarely falls. But the
summertemperatureat HumboldtBay is well knownto be much
coolerthan that of even somewhathigherregionsin the interior,
up to an altitude of at least 4,000 or 5,000 feet,because of the
eastwardmovingair-currents, which are coolestwherethey first
leave the sea surface and warm up as theypass fartherand far-
therinland. We are thereforeled directlyto the finalinference
that the summertemperatureat sea level about Humboldt Bay

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120 THE AMERICAN NATURALIST [VOL. LI

closely approximatesthe summertemperatureat from6,000 to

9,000 feet on Mount Shasta and above 7,000 feet on the Warner
Mountains. In these three areas, the air is cooler in summer
than in the interlyingareas and thus betteradapted to the finely
adjusted requirementsof the Oregon jay. Summer tempera-
ture, betweencertain degrees,is one critical factor.
Three more factors present themselvesfor considerationin
connectionwith the Oregon jay, those of humidity,rainfall and
cloudiness. HumboldtBay lies in the mosthumid and continu-
ously rainy section of California. Mount Shasta and the War-
ner Mountains are relatively arid, the latter most notably so.
It would appear, therefore,that humidity,rainfall and cloudi-
ness had little or nothingto do with cuttingoffthe range of this
bird, thoughone or otherof these factorsmay have been respon-
sible for the very slightly darker tone of color which distin-
guishesthe coast jays (subspeciesPerisoreubsobscurus obsetrus)
fromthose in the interior (P. o. gr'isems). But, however this
may be, it is clear that temperaturemust dominategreatlyover
the three factorsnamed in checkingdissemination.
In summary,we may thereforedispose of the followingfac-
tors as havinglittleor no effecton the distributionof the Oregon
jay as a species: the nature or quantity of its food supply, at
mosphericdensity and pressure, cloudiness,rainfall, humidity
of the air or soil, and wintertemperature. This eliminatesall
but the two factors: shelter of a sort provided by the forest
habitat, and temperatureof the summerseason.

THE CASE OF THE CONY

The cony or pika is a mammal representedin California by

fourquite similar races (Ochotona taytori,0. schisticepsschisti-
in
ceps, 0. s. Muiri, and 0. s. albatus), whichagreeclistributionally
occupyinga very restrictedhabitat along high mountaincrests.
I know of no place in central California where conies range
below an altitude of about 8,000 feet,and they range upwards
to fully 12,000 feet in the vicinityof Mount Lyell. They thus
occupy an altitudinal belt between extremes4,000 feet apart.
With regard to zones of vegetationconies live fromconsiderably
below timberlineto considerablyabove timberline. Extended
observationshows that their existence is in no way correlated
withthat of treesor shrubsof any sort. Like theirrelatives,the
rabbits,they feed entirelyon low vegetation,biennials mostly;

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No. 602] SHORTER ARTICLES AND DISCUSSIONS 121

but unlike mostkinds of rabbitsthey are strictlydependentfor

safety from enemies upon rocks, especially where these are
looselypiled as in talus slopes and so afforddeep retreatswithin
their interstices. The whole equipment of a rabbit is clearly
adapted to foragingin the open, its keen hearing and eyesight
quickly warning it of the approach of enemies,and giving it
time to escape by means of its unusual running powers. But
the cony is equipped in a verydifferent way, as it has relatively
small ears and eyes, and small hind legs. It is compelled to
forage close to or beneath cover. In fact in fieldobservationsit
is rarely seen on the move except momentarily,and then only
between or beneath angular granite blocks,where it grazes on
such little patches of vegetationas are withinimmediatereach.
It is clear fromnumerousobservationsthat the conyis sharply
restrictedin a large part of its range by the rock-pilehabitat.
Even at favorable altitudes it is not found away from this
refuge. There are obviously,however,one or more additional
factorsin its distribution. In many parts of the Sierras, talus
slopes occur fromnear the highestsummitsdown to the foothills.
As examples of these,one may cite the vast earthquake taluses
of the YosemiteValley proper,which occur almost continuously
down to and below the 4,000-footcontour. These taluses have
been searched diligentlyboth by trapping and hunting,without
our naturalistsfindinga trace of conies below 8,000 feet. The
animals are easy to detect,by reason of their characteristiccry,
utteredat any time duringthe day, thoughmoreparticularlyin
the morningand the evening,and by the accumulationsof their
feces,the pellets constitutingwhich are, in size, shape and tex-
ture,unlike those of any othermammal. What is it, then,that
limitsthe conies downwardon the westernflankof the Sierras,
wheretheirnecessaryrockhabitat is continuous,and wherefood
of the rightsort is also continuous? Let us trybarometricpres-
sure, and atmosphericdensity,which may properly be consid-
ered together. These conditionschange sensibly with altitude
and, if we take into account California alone, the facts would
seem to entitlethemto serious considerationas active delimitors
of the coniesdownward. But as we trace the range of the conies
far to the northwardwe are led to a differentconclusion. The
altitudinallimits of theirrange is found to descend quite regu-
larly towardsthe north,until, in the case of one race, even sea
level is reached,at Bering Sea. Clearly,conies,generically,are

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122 THE AMERICAN NATURALIST [VoL. LI

thus proven not to be affectedby atmosphericpressure,or by

atmosphericdensity,at least in as far as it is modifiedby alti-
tudes up to 12,000 feet. The same fact-depression of range
towards the north-discloses a third concomitantof altitude,
which is also a concomitantof latitude, namely, temperature,
and this is beyond doubt the determiningfactor. As the iso-
thermsdip toward sea level to the northwardso does the range
of the genus'Ochotona. We have, therefore,by study of geo-
graphical distributionin this case establishedtwo importantcon-
trollingfactors,namely (1) safetyrefugesof a sort providedby
talus slopes and glacial moraines; (2) temperature,at least
downwardbelow the degree,correlatedin the mountainsof Cali-
fornia by a mean annual or summercomputationor forabriefer
period at the time of reproduction,with an altitude of eight to
twelve thousand feet,according to latitude, slope exposure and
air currents.
It is not possible for one to say fromthe data in hand what
the directcontrollingfactorsof the upward limitsof the cony's
range may be. Taluses extendup to the highestpeaks,but there
is no growthof grass above about the 12,000-footcontoureven on
the mostfavorableslopes. As the disappearance of the coimy in
the higher altitudes is coincidentwith the disappearance of its
food, it appears as if failure of food alone were the delimitor
here; but we have no way of showingthat even if food did con-
tinue the cony would be restrictedupward, as it certainly is
downward,by a change in temperaturebeyond some critical
point. The cause of its delimitationdownward,however,re-
mains clear.
THE CASE OF THE Rosy FINCH
In the case of the bird called genericallyLentcosticte,or rosy
finch,we find-a condition astonishinglysimilar to that of the
cony. In fact almost the entire preceding account could be
made relevant'here, by merelysubstitutingthe term rosy finch
for cony. The ranges, altitudinal and geographical,of the two
animals are almost identical. The only obvious differencesap-
pear in their ecologic relations, and consist in the lesser de-
pendence of the bird upon shelterand in the dissimilarnature
of its food. The rosy finchforages gregariouslyon the open
slopes, near timberlineand above, though its nest is hidden
away in the cleftsof rockledges and taluses. It shuns the trees

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No. 602] SHORTER ARTICLES AND DISCUSSIONS 123

and bushes even whereit ranges well below timberline. It feeds

winterand summerupon seeds of dwarfedvegetation,including
those of grass and herbs of various sorts. As far as I can see,
its food and feedinghabits are identicalwith thoseof such other
fringillidsas goldfinchesand siskins. Yet the leucosticte,by
the same testsas were used with the cony,is beyond any conten-
tion limiteddownwardby an increase of temperature. We find
the bird to possessvarious adaptive featuresin commonwith cer-
tain arctic finches,such as tuftsof bristle-likefeathersover the
nostrilsto prevent fine snow fromentering. These enable the
bird to spend the long winteron the cold wind-sweptridges,but
at the same time would hardly preventthe bird's dropping to
warmerclimes if the heat were not a stronglydeterrentfactor.
Cases of coincidence,as instanced by that of the cony and
leucosticte,among animals of widely different powers of locomo-
tion and ecologic position,are the rule, not the exception,and
of the controllingfac-
impel the observerto belief in the efficacy
tor above mentioned.

THE CASE OF THE REDWOOD CHIPMUNK

The redwood chipmunk (Eutarnias townsendiochrogenys)is

an animal confinedto a very narrow but exceedinglylong dis-
tributionalarea extendingsouth fromthe Oregon line as far as
Freestone,Sonoma County. Throughoutthis belt it is conspicu-
ously numerous,and is usually the only species of chipmunk
present,so that the limits of its range have been easy to mark
definitelyalong the several lines explored. This rodent, by
various geographic tests similar to those I have recounted for
other birds and mammals,is clearly delimited away from the
coast at the bounds of the well-knownfog-beltto whichthe red-
wood tree and numerousotherplants as well as animals belong.
The chipmunk,however,depends in no way upon the redwood
or any other one plant species as far as I can see, but feeds
upon a great variety of seeds and fruits,like many of its con-
generselsewhere.
That temperatureis also a delimitingfactoris shownin parts
of therange of the redwoodchipmunk. But atmospherichumid-
ity or cloudiness or rainfall, factorswhich I have in this case
failed to dissociate, together constitute or include the chief
controls.

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124 THE AMERICAN NATURALIST [VOL. LI

THE CASE OF THE BELTED KINGFISHER

It is to be observedthat specialization for gettinga particu-

lar kind of food invariablybringswith it restrictionof range to
the territoryproviding that kind of food. The northwestern,
belted kingfisher(Ceryle alcyon caurina) is a good example of
.this. In California we find this bird present at various times
of theyear bothalong theseacoast and along variousfish-support-
ing streams,fromthe Colorado River to the Klamath River and
up the mountainstreamsto at least as high an altitudeas Yosem-
ite Valley. The kingfisheris seen during migrationin many
places away fromstreams,but it tarriesat such timesonlywhere
its natural diet can be procured,as, on occasion,at fishponds in
city parks. There is a unique instance of a kingfisherobserved
on the desert catching lizards, but exceptional occurrencesof
this kind are of course not to be given considerationin making
generalizations.
It is observablefurtherin regard to this species of kingfisher,
that it must have earth banks in which to excavate its breeding
tunnels. Lack of these along any stream,otherwisefavorable,
preventsthe bird fromstaying there throughthe season of re-
production. Furthermore,thereis also obvioustemperaturere-
striction;for,given a fish-producing stream,with banks appar-
entlywell suited for excavationof nestingplaces, such as is the
Colorado River and its distributaries,and the summertempera-
ture mustbe at least below that of southernCaliforniasouth of
the 35th parallel. That all such streamsare well supplied with
kingfishers in winter,and are forsakenonly duringthe hot sum-
mer,seemsto showthat a relativelycool temperatureis forthem
in some way or anotheressentialto successfulreproduction.
We find,then,in the case of the beltedkingfisher, that the fac-
tors of a requisitekind of food, and a requisitekind of nesting
place, both having to do with the structuralpowers and limita-
tions of the bird, togetherwith the factorof the temperatureof
the summerseason, are those that account for the distribution
of the species withinthe state of California,as we findit.

THE CASE OF THE MEADOWLARK

The western meadowlark (Sturnella neglect) is a bird of

relativelyomnivorousdiet. Note that I say relatively,for the
word omnivorousunmodifiedwould apply only to such an ani-

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No. 602] SHORTER ARTICLES AND DISCUSSIONS 125
mal as would eat the sort of food that any animal eats, and this
is an obvious impossibilityfor the meadowlarkwhen we con-
sider such uncommonarticles of diet as wood and petroleum.
Compared with many other birds, the meadowlarkdoes use as
food a very wide range of plant and animal objects. This food,
however,is restrictedto a particular habitat source, namely to
the meadow. The bird's entire equipment specializes it for
successful food-gettingand for escape from enemies upon a
grassy plain or meadow. And it is a matterof commonobser-
vation that its range is sharply delimitedin most directionsat
the margin of the meadow habitat, as where this is interrupted
by forest,brushland,marsh,rock surface or sand flat. This is
a conspicuousexample of what we may call associationalrestric-
tion. But it is not the only way in which the meadowlarkis
hemmedin. In this connectionCalifornia again provides crit-
ical distributionalevidence.
We findmeadowlarksoccupyingpracticallyeveryappropriate
meadow,large and small, fromthe Mexican line to the Oregon
line and fromthe shores of the Pacific to the Nevada line, ex-
cept above a certainlevel on the higher mountains. In travel-
ing up the west flankof the Sierras, and this I have now verified
along three sections,meadowlarkscease to be observed at ap-
proximatelythe 4,500-footlevel, and this in spite of the factthat
above that altitude meadowsare found which are to all appear-
ances ideal for meadowlarkrequirements. I need only referto
such seeminglyperfect summerhabitats as Monache Meadows
and Tuolumne Meadows. And though, in the winter these
would be uninhabitable,so are othermeadows (as those in the
Modoc region,for instance), which are in summerwarm and at
that season abundantlyinhabitedby meadowlarks. By the elim-
ination then upon proper grounds of various factorsfromthe
list, we have left only three possible factorsin this upward de-
limitation,namely, decreased atmosphericpressure, decreased
air density and decreased temperatureof the summerseason.
Since meadowlarksexistat correspondingaltitudesin thewarmer
thoughelevated Great Basin region,and since it has been pos-
sible to eliminatepositivelyand in a similar way the firsttwo
factorsin the cases of many otherbirds and mammals,these fac-
tors are presumablywithoutinfluenceon the meadowlark; and
thereis leftbut one-temperature.
Within the state of California, meadowlarks,without the

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126 THE AM1ERICAN NATURALIST [VOL. LI

slightestdetectable subspecificmodification,thrive under both

the cloudy, humid conditionsof the northwestcoast belt and
under the relativelycloudless,arid conditionsof Owens Valley.
Factors of humidity,of air and soil, cloudiness,and light in-
tensity,seem to avail nothingin checkingtheir spread. With
such a degree of associational specialization as is exhibitedby
thesebirds thereis little chance of a serious competitivestruggle
with other vertebrates,and no evidence of such has been ob-
served. As far as California is concerned,the' meadowlark's
range is thus only limited associationallyand zonally, that is
by the extent of its particular meadow habitat and by dimin-
ished summertemperaturebelow some critical point.
The meadowlarkwell illustratessome furtherfacts with re-
gard to distribution. In Calfiforniait is unquestionablyon the
increase as regards total population. This is due chieflyto the
great extensionof habitable territoryresultingfromman's oc-
cupancy and cultivationof the land, bare plains, brushlandsand
even woods being replaced by irrigatedalfalfa and grain fields.
These the meadowlarksfindsuitable and invade,because of their
expansive reproductivity,and soon populate to the fullest ex-
tent permittedby the minimumannual food supply. In other
words, associational barriers have moved, to the advantage of
this particular bird, though at the same time to the disadvan-
tage of endemicspecies of different predilections. I should esti-
mate that the total meadowlarkpopulation in the San Joaquin-
Sacramento basin is now fully three times what it was thirty
years ago.
Animal distributionis not fixed. It changeswith the shifting
of the various sortsof barriers,and doubtlessalso as a result of
a gradual acquisitionby the animals themselvesof the power to
overstepbarriers,as by becominginured to greateror lesser de-
gree of temperature. The power of such accommodation,or in-
herent plasticity,evidentlyvaries greatly among differentani-
mals; and at best its operationis very slow. Many species have
proved stubbornand have been exterminated,as the factor-lines,
or barriers,shifted. By the shiftingof, say, two criticalfactor-
lines towards one another,the existenceof a species may have
been cut offas by a pair of shears.

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No. 602] SHORTER ARTICLES AND DISCUSSIONS 127

SUMMARY

In this paper I have enumeratedvarious factors thoughtto

be concernedwith the controlof the distributionof vertebrate
animals. A numberof birds and mammalshave been cited to
showhow we mayuse our moreor less detailedknowledgeof their
ranges so as to demonstratethe operation of one or several out
of the many possible factors as limitersto distribution. The
methodemployedis one of examination,comparisonand elim-
ination, applied to all parts of the margin of animals' ranges.
The range of any one animal must be examinedat all points of
its peripheryin order that all of the factorsconcernedmay be
detected. One factormay constitutethe barrier in one section
of the peripheryof the range of a species,a totallydifferentfac-
tor in another section.
The results of the geometric ratio of reproductionwould
bring about areas of occupancy in the shape of perfectcircles.
But we neverfindsuch symmetricalranges. The very fact that
the outlinesof the ranges of animals are extremelyirregularis
significantof the criticalnature or inexorablenessof the factors
which delimitthem. These factorshave to do with the evolu-
tion, persistenceand exterminationof species.
Note that we always have to take into account,in attempting
to discernfactorsof limitation,the animal's own inherentstruc-
tural equipment. This prescribesrestrictionat once in certain
regards. Referringagain to our list of suggestedfactors,we find
the long-emphasizedones of land to aquatic species and bodies
of water to terrestrialspecies really presentingan extrememani-
festationof associational restriction. Food source, methodsof
food-getting and safetyrefugesare involved.
It is to be noted furtherthat the factorsare various and that
the most importantfactor for one species may prove of little
effectwith another species. Species do not react uniformlyto
the same environment. It is undoubtedlyalways a combination
of factorswhich accounts for an animal's geographicrange in
all parts of the peripheryof that range. It is most certainly
never one factoralone. No one will claim that temperatureis
the only delimitingagent in controllingvertebratedistribution;
nor could this claim be made for humidityalone, or for food
supply alone, or for safety of breeding-placesalone.
Given a large continuousarea, however,as upon the North

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128 THE AMERICAN NATURALIST [VOL.LI

American continent,one single factor does happen to loom up

as being the most frequentdelimiterof distribution,or even the
ultimatelyeffectiveone, in greater or less degree, even though
other factors be effectivealso. This factor is temperature.
The cases cited illustratethe tenetthat in some directionor an-
other,temperaturebeyond certain limits,up or down, cuts off
furtherdissemination. This is part of the basis of the life-zone
idea. But, as I have tried to bring out above, this fact is in no
way antagonisticto the claim that otherfactors,as of humidity,
food supply, and shelter,also figure critically,giving a basis
for recognizingfaunal areas and associations. Finally, if our
discussionof the subject has been sound, it is evidentthat data
secured throughfieldobservationcan be so employedas to bring
results essentiallysimilar to, and as conclusiveas, those secured
throughlaboratoryexperimentation.
JOSEPH GRINNELL
MUSEUM OF VERTEBRATEZOOLOGY,
UNIVERSITYOF CALIFORNIA,
November 1, 1916

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