Grinnell 1917 the Niche-relationship of the California Thrasher

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    VI'ilXyXIV GRINNELL,heCaliforniahrasher. 427

    into the body of the partly eaten bantam and replaced t in the

    same spot where he found it. Next morningthe seemingly m-

    possible as madea practicalcertainty, or he foundthe body of a

    screech wl with the clawsof one foot firlnly imbedded n the body

    of the bantam. He very kindly presentedme with the owl which,

    upon dissection, roved to be a female, ts stomacheontalninga

    very considerablemountof bantam fleshand feathers, ogether

    with a great deal of wheat. (It seelnsprobable hat the wheat

    was accidentallyswallowedwith the crop of the bantam during

    the feast,but therewas so much hat it seelns trange he owl did

    not discard t while eating). How a bird only 9.12 inchesn length

    could have dealt out such havoc in so short a time is almost in-

    credible,but, althoughpurely circumstantial,he evldenee gainst

    the owl appeared ltogether oo strong or evena reasonable oubt.

    The doctorand I wished o make as certain as possible, owever,

    so the poisonedbantam was replacedand left for severaldays,

    but without any further results. For the abovementioned easons

    I am rather doubtful as to the net value of this owl from an

    economictandpoint,lthoughirdsn a wildstatwould ot

    give theln such opportunities or such wanton killing as birds

    enclosedn pens.

    THE NICHE-RELATIONSHIPS OF THE CALIFORNIA

    THRASHER?

    BY JOSEH GRINNELL.

    TUE California Thrasher (Toxostomaedivivum) s one of the

    severalistinctird ypes hichharacterizeheso-calledCali-

    fornian Fauna. Its range is notably restricted, even more so

    than that of the Wren-Tit. Only at the south does he California

    Thrasher occurbeyond he limits of the state of California, and in

    that directiononly as far as the San PedroMartit Mountainsand

    Contribution from the Museum of Vertebrate Zo51ogyof the University of California.

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    [Auk

    428 GRINNELL,heCaliforniahrasher. toct.

    San Quintin, not more than one hundredand sixty milesbelow he

    Mexican line in Lower California.

    An explanationof this restricteddistribution s probably o be

    found n the closeadjustmentof the bird in variousphysiological

    and psychologicalespects o a narrow range of environmental

    conditions. The nature of these critical conditions is to be learned

    throughan examination f the bird's habita.t is desirableo

    make suchexamination t as many points n the general angeof

    the species s possible ith the objectof determininghe elements

    common o all thesepoints,and of these he onesnot in evidence

    beyond the limits of the bird's range. The followingstatements

    in this regardare summarizedrom the writer's personal xperience

    combinedwith all the pertinent nformationafforded n literature.

    The distributionof the CaliforniaThrasheras regards ife-zone

    is unmistakable. Both asobservedocallyand over ts entire ange

    the species hows loseadherenceo the Upper Sonorandivision

    of the Austral zone. Especiallyupwards, s it always sharply

    defined. For example, n approachinghe sea-coast orth of San

    FranciscoBay, in SonomaCounty, where the vegetation s pre-

    vailingly Transition, thrashersare found only in the Sonoran

    islands, namely southerly-facing ill slopes,where the maxi-

    mum insolationmanifests ts effects in a distinctive chaparral

    containing uch ower zoneplants as Adenostoma. Again, around

    Monterey, to find thrashers ne must seek the warm hill-slopes

    back from the coastalbelt of conifers. Everywhere have been,

    the thrashers eem o be very particularnot to ventureeven a few

    rods into Transition, whether the latter consistof conifersor of

    high-zonespeciesof manzanita and deer brush, though the latter

    growth resembles losely n densityand general appearancehe

    Upper Sonoranchaparraladjacent.

    While sharplydelimited,as an invariable ule, at the upperedge

    of UpperSonoran,he CaliforniaThrashers not socloselyestricted

    at the loweredge of this zone. Locally, individuals occur, and

    numbersmay do so where assoeiationalactors favor, down well

    into Lower Sonoran. Instancesof this are particularlynumerous

    in the San Diegan district; for example, n the Lower Sonoran

    washes at the mouthsof the canyons longthe southbaseof the

    San Gabriel Mountains, as near San Fernando, Pasadena, and

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    ol. XXXI1

    1917 J GRIXnL, The California Thrasher. 429

    Azusa.A noticeablehing n thisconnection,owever,s that,

    on the desert lopes f the mountains, hereToxostomaecontei

    occursn the desertlooras an assoeiationalomologuef T.

    redivivumn theLower onoranone,he atter staysut ar

    3. Toostorna. pasadenense

    ' - ' Specimensxamined

    Publishedecords

    Figure 1.

    moredosely; that is, it straysbut little or not at all below he

    typicalconfinesf its ownzone,namelyUpperSonoran.The

    writer'sieldwork n thevicinity f Walker ass, ernCounty,

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    [-Auk

    430 GRINNELL,heCaliforniahrasher. LOct.

    provides ood llustrations f this. A tongueor belt of Lower

    Sonoran extends from the 3/ohave Desert over the low axial moun-

    tain ridgeat the head of Kelso Creek and thencedownalong the

    valley of the South Fork of the Kern River nearly to Isabella.

    Leeonte's Thrasher is a conspicuouslement in this Lower

    Sonoran invasion, but no California Thrashers were met with in

    this regionbelow he belt of goodUpper Sonoran n the flanking

    mountain ides, s markedby the presencef diggerpine,blueoak,

    sumaeh, silk-tasselbush, and other good zone-plants. Similar

    zonal relationships re on record from San GorgonioPass, River-

    sideCounty,as well as elsewhere.

    Reference owto' he general angeof the bird underconsidera-

    tion (seep. 429), as comparedwith a llfe-zonemap of California

    (PacificCoastAvifaunaNo. 11, Pls. I, II), will show o a remas'k-

    abledegree owclosely he formercoincides ith the UpperSono-

    ran zone. The thrasher s, to be sure,one of the elementsupon the

    presencef which his zonewasmarkedon the map; but it was

    only one of many, both plant and animal; and it is concordance

    wi];hheaggregatehat ssignificant. iagnosisfzonationimi-

    larly is possiblen scores f placeswherechangen altitude(which

    as a rule means hangen temperature)s the obviousactor,as up

    the west flank of the Sierra Nevada, or the north wall of the South

    Fork valley, alreadyreferred o, in Kern County, or on the north

    wall of the San Jaeinto 3/ountains. The California Thrasher is

    unquestionablyelimited n its range n ultimate analysis y tem-

    peratureeonditlons.The isothermie rea it occupiess in zonal

    pax'lance, pper Sonoran.

    The secondorder of restriction s faunal, using this term in its

    narrowed ense,ndleatingdependenceponatmospheric umidity.

    The CaliforniaThrasherdoesnot range nteriorly nto excessively

    arid country,although he UpperSonoran onemay, as around he

    southern end of the Sierra Nevada, continue uninterruptedly

    towardsthe interior in a generally atitudinal dh'eetion. This

    is true where extensiveareas are considered, ut locally, as with

    zones, ndividualsor descent-lines ay have invadedshort dis-

    tanees beyond the normally preferred conditions. An example

    of this situation s to be foundon the north and westslopes f the

    San Jacinto3/ountains,where CaliforniaThrashers ange around

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    Vol. XXXiV1

    1017 J GRINNELL,heCaliforniahrasher. 431

    onto arid chaparralslopes,nterminglingwith such arid Upper

    Sonoranbirds as Scott'sOrioleand the Gray Vireo. It is question-

    able, however, as to what extent faunal restriction really operates

    in this case; for reference o the zone map, again, shows hat a

    vast tract of LowerSonoran,ying to the eastof the desertdivides,

    extends ontinuously orth to the head of OwensValley. Really

    the onlyunbroken ridgeof UpperSonoranowards he east rom

    the west-Sierran habitat of Toxostoraa redivivura is around the

    southern ndof the SierraNevada - a very narrowand ongroute

    of possible migration,with consequentactors unfavorable o

    invasion, rrespective f either temperatureor humidity, such as

    interruptedassociationalequirements nd small aggregate rea.

    In this particularbird, therefore, aunalrestrictionmay be of minor

    importance,as comparedwith zonal and associationat ontrols.

    That faunal conditions ave had their influenceon the species,

    however, s shownby the fact of geographic ariationwithin its

    range. The thrasher throughout ts habitat-as-a-whole,s sub-

    jected to different degrees f humidity. Amount of rainfall is,

    in a generalway, an index of atmospheric umidity, thoughnot

    withoutconspicuousxceptions. Comparinghe mapof the ranges

    of the subspeciesf T. redivivurap. 429) with a climaticmapof the

    State, direct concordances observedbetween areas of stated rain-

    fall on the latter and the rangesof the respective ubspecies. t

    will be seen that the race T. r. pasadenenseccupiesan area of

    relatively ow humidity, he race T. r. sonoratef higherhumidity

    and the race T. r. redivivura f highesthumidity, in fact a portion

    of California's og-belt. The distinctivecolor-tones eveloped re,

    respectively,of gray, slate and brown casts. In the thrasher,

    therefore,we may look to faunat nfluences s havingmost to do

    with differentiationwithin the species. In this case t is the faunal

    variationover the occupied ountrywhich s apparently esponsible

    for the intra-specificudding, r, in otherwords, he origination f

    new specific ivarications.

    qerever it occurs, nd in whichever f the three subspeciest is

    represented,he CaliforniaThrasherevinces' trongassociational

    predilections. t is a characteristiclement n California'samous

    chaparralbelt. Where this belt is broadestand best developed,

    as in the San Diegandistrictand in the foothill regions ordering

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    [Auk

    432 GRINNELL,heCaliforniahrasher. [Oct.

    the great interior valleys, there the Thrasher abounds. The

    writer's personal ield acquaintancewith this bird givesbasis or

    the followinganalysisof habitat relations.

    The California Thrasher is a habitual forager beneath dense

    and continuous over. Furthermore, probably two-thirdsof its

    foraging s done on the ground. In seeking ood aboveground,

    as when patronizingcascarabushes, he thrasher arely mounts

    to an exposedposition,but only goesas high as is essential o

    securing he coveted ruits. The bird may be characterized s

    semi-terrestrial,but always dependentupon vegetationalcover;

    and this covermust be of the chaparral ype, open next to the

    ground,with stronglynterlacing ranch-work nd evergreeneafy

    canopy lose bove- not orestunder-growth,r close-set,pright

    stems as in new-growthwillow, or matted leafage as in rank-

    growing nnualherbage.

    The Thrasher s relativelyomnivorousn its diet. Beal (Bio-

    logicalSurveyBulletinno. 30, p. 55) examined 2 stomachs f

    Toxostomaedivivum nd found hat 59 percentof the foodwasof a

    vegetable ature and 41 animal. A large part of this food con-

    sistedof ground-beetles,nts,andseeds, uchas are undoubtedly

    obtainedby workingover he litter beneath haparral. The bird's

    mostconspicuoustructural eature, he longcurvedbill, is used o

    whiskaside he litter, and also o dig, pick-fashion,nto softearth

    where nsectsie concealed. Groundmuch requented y Thrashers

    shows umerousittle pits in the sollsurface,ess han an inchdeep,

    steepon onesideand with a little heapof earthpiledup on the

    oppositeide. Asalreadyntimated,heThrasher t timesascends

    to thefoliage bove,or fruit anddoubtlessomensects. Much n

    thewayof berries ndseedsmayalsoberecoveredrom he ground

    in what s evidently he Thrasher's wnspecialized ethodof food-

    getting. Evengrantinghisspecialization,do not seewhy the

    chaparral,lone, hould fford he exclusiveorage-ground;or the

    samemodeof food-gettingught o be just as usefulon the forest

    floor,or evenon the meadow. The furtherfact, of widelyomni-

    vorousdiet, leadsoneto concludehat it is not any peculiarityof

    food-source,r way of getting t it, that alone imits he Thrasher

    associationally.We must look farther.

    The amateurobserver, r collector f specimens,s struckby the

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    'Vol. XXXI]

    1917 J GRXNNLL, he CaliforniaThrasher.

    433

    apparent shyness of the Thrasher--by the easewith which it

    eludescloseobservation,or, if thoroughlyalarmed, escapes etec-

    tion altogether. For this protective effect the bird is dependent

    upon appropriatecover, the chaparral, and upon its ability to

    co-operaten makinguse of this cover. The Thrasherhas strong

    feet and legs,and muscular highs,an equipmentwhichbetokens

    powersfrunning;he &il sconspicuouslyong, s n manyun-

    ning birds; and correlatlvely he wings are short, rounded,and

    soft-leathered, ndicating little use of the flight function. The

    colorsof the bird are non-conspicuous-blended,dark and light

    browns. The nests of the Thrasher are located in dense masses

    of foliage, rom two to six feet above he ground, n busheswhich

    are usuallya part of its typicalchaparralhabitat. In only excep-

    tional casess the chosen estingsite located n a bushor scrubby

    tree, isolatedmoreor less rom the main body of the chaparral.

    Thesevariouscircumstances,hichemphasize ependencepon

    cover, and adaptation in physical structure and temperament

    thereto,go to demonstratehe nature of the ultimate associational

    niche occupiedby the California Thrasher. This is one of the

    minor nicheswhichwith their occupantsll togethermake up the

    chaparral association. t is, of course,axiomatic that no two

    species egularly establishedn a single auna have precisely he

    same iche elatlonships.

    As a final statementwith regard to the CaliforniaThrasher,

    we may conclude,hen, that its range s determined y a narrow

    phaseof conditions btainingn the Chaparralassociation,ithin

    the California auna, and within the Upper Sonoran ife-zone.