THE AUK

A QUARTERLY JOURNAL OF
ORNITHOLOGY

VOL. 104 APRIL 1987 NO. 2

EFFECTS OF NESTLING DIET ON GROWTH AND

ADULT SIZE OF ZEBRA FINCHES
(POEPHILA GUTTATA )

PETER T. BOAG
Departmentof Biology,Queen'sUniversity,Kingston,OntarioK7L 3N6, Canada

Al•STRACT.--Manipulation of the diet of Zebra Finch (Poephilaguttata) nestlings in the

laboratoryshowedthat a low-quality diet reducedgrowth ratesof nine externalmorpholog-
ical characters,while a high-quality diet increasedgrowth rates.The growth of plumage
characterswas least affectedby diet, while growth ratesof tarsusand masswere most af-
fected. The treatmentsalso produced differencesin the adult size of experimental birds,
differencesnot evident in either their parentsor their own offspring.Diet quality had the
strongestimpact on adult massand tarsuslength, while plumageand beak measurements
were lessaffected. Analysis using principal componentsand characterratios showed that
the shapeof experimentalbirds was affectedby the experimentaldiets,but to a minor extent
comparedwith changesin overall size. Significantshapechangesinvolved ratiosbetween
fast- and slow-growingcharacters.The ratiosof charactersthat grow at similar, slow rates
(e.g. beak shape) were not affectedby the diets. Environmental sourcesof morphological
variation should not be neglectedin studiesof phenotypicvariation in birds. Received5 June
1986,accepted30 October1986.

MORPHOLOGICAL differences between indi- fitness, and weather was seen in the nonran-
vidual birds are often assignedfunctional sig- dom survival of House Sparrowscollectedby
nificance, whether those individuals are of dif- Hiram Bumpus following a winter storm
ferent species,different sexes,or different-size (O'Donald 1973, Fleischer and Johnston 1982).
members of the same sex (Hamilton 1961, Se- Recently, investigators have tried to dem-
lander 1966,Clark 1979,James1982).In grani- onstrate a genetic basis for intraspecific mor-
vores such as Darwin's finches(Geospiza spp.), phological variation in birds. Several studies
morphologicaldifferenceshave been related to have shown that variation in morphological
variation in feeding efficiency at all three of traits of ecologicalinterest is heritable (seere-
these levels. Differences in feeding efficiency view by Boag and van Noordwijk 1986). Al-
have in turn been correlated with variation in though such analysesare difficult to perform
fitness due to spatial or temporal variation in and interpret, they have provided one of the
seed availability (Boagand Grant 1981, Schlu- few meansavailable to test the plausibility of
ter and Grant 1984). Intraspecific morphologi- the hypothesisthat natural selectionis the pri-
cal variation in several species,such as House mary determinant of the array of phenotypes
Sparrows (Passerdomesticus; Johnstonand Se- maintained within avian populations.
lander 1971), Red-winged Blackbirds(Agelaius One common misunderstandingof heritabil-
phoeniceus; Power 1969),Great Tits (Parusmajor; ity studies is that even if variation in a trait
Hamilton 1961), and others (James 1970) has withintwo different populationsis highly her-
been correlated with clinal variation in cli- itable, this does not guaranteethat the differ-
mate. A classicexample of the correlation be- ence in the characterbetweenthe populations
tween intraspecific morphological variation, has a geneticbasis.For instance,regressionsof
155 The Auk 104: 155-166.April 1987
156
offspring bill size on parental bill size were
identical and significant in two years of data
collectedfor Geospiza fortis(Boag1983).This in-
dicated that the variation was highly heritable
in both years. However, y-intercepts in these
sameregressionsdiffered significantly(seeBoag
1983:fig. 1). Boag(1983) hypothesizedthat dif-
ferent population densitiesand territory sizes
in the two yearsled to differencesin the quan-
tity or quality of food fed to young, with the
result that young grew to different adult sizes.
Although it hasbeen known for sometime that
nestling diets can alter growth rates (Bryant
1978), few authors have suggestedthat varia-
tion in growth rate contributesto adult mor-
phologicalvariation in wild birds (but seeSmith
and Zach 1979; James 1982, 1983; Boag 1983;
Alatalo and Lundberg 1986). Laboratory data
on birds (Allee and Lutherman 1940, Johnson
1971) and mammals (Harrison 1959, Lister and
McCance 1965) suggest that diet in early life
can affect adult size.
This possibility is supported by elegant ex-
periments on Red-winged Blackbirds (James
1983). Reciprocaltransplants of Red-wings be-
tween either end of two separate geographic
clines in body size and shapeshowed that the
transferredyoung grew to resembletheir foster
parents more than their biological parents.
Jamesconcludedthat Red-wing clines in mor-
phology had a large nongenetic component,
and that there probably existed complex co-
variation between genetic and environmental
causesof individual differences in morpholo-
gy.
The heritability studiesdiscussedabove, as
well asthe transplantexperiments(James1983),
signal a growing need to considerthe origins
of phenotypic variation in avian populations.
Suchawarenesscameearly to ecologistswork-
ing with other taxa.For instance,the morpho-
logical plasticityof plants routinely forcedbot-
aniststo dissectecotypicvariation in their study
populations before invoking adaptive argu-
ments concerningindividual differences.This
approachwas epitomizedby the classicecotype
transfers conducted by Clausen et al. (1940).
Until recently, most avian ecologistshave as-
sumedimplicitly that birds have essentiallyde-
terminate growth, and that phenotypicdiffer-
encesbetween adults accuratelyreflect genetic
differences, which are amenable to evolution
by •,atural selection.
PETER
J. BOAG [Auk,Vol. 104
I report a simpleexperimentwith laboratory-
raised Zebra Finches (Poephilaguttata).I show
that qualitativedifferencesin the diets fed by
parentsto their young producesubstantialdif-
ferences in the growth rates of several mor-
phologicalcharacters.When these young are
followed until adulthood, the different nest-
ling diets produceadult birds with significant-
ly different morphologies.
METHODS
Adult Zebra Finches were selected at random from
an outbred laboratorycolony and mated to produce
14 pairs. Eachpair was placed in a 50 x 30 x 40 cm
breeding cage, provided with a wicker nest cup,
shreddedburlap nestingmaterial,cuttlebone,an oys-
ter shell-grit mixture, and water. Cageswere main-
tained indoors on a 14L:10D lighting schedule at
constanttemperatureand humidity, visually but not
acousticallyisolatedfrom eachother. Cageswere as-
signedat randomto one of three experimentalgroups,
which were treated identically exceptfor the quality
of food available from the end of incubation until
the youngwere approximately14 weeksold. Parents
were removed when young were about 35 days of
age. Breeding under all treatments commencedsi-
multaneously.
Families in the "standard-quality diet" treatment
groupwere fed finch seedmix (red and white millet,
with canary, rape, and flax seed) and standardnest-
ling food mixture (see Table 1) ad libitum.The seed
mix contained about 14% crude protein, while the
nestling mix had about 24% protein, basedon Kjel-
dahl analyses(Skoog and West 1969). In addition,
thesefamiliesreceivedminced,hard-boiledhen's egg
with added multivitamin-mineral powder (Perve-
line•, 8 in 1 ProductsInc., about 1 g/100 g of egg)
oncea week (about 1 g wet massegg mixture for each
adult or young in cage). Three days after the egg
ration, eachcageof birds receivedapproximately5 g
wet massof fresh lettuce or spinach.The four stan-
dard-dietpairseachproducedone brood,with a total
of 6 male and 5 female offspringsurviving to 1 yr of
age or more.
Families in the "high-quality diet" group were
given the standardseedmixture,and a high-protein
nestling mixture (approximately38%protein; seeTa-
ble 1) ad libitum.They receivedabout 1 g per bird of
the egg mixture every day and 5 g of fresh greens
every other day. Five high-diet pairs eachproduced
one brood, for a total of 10 male and 4 female off-
spring at 1 yr of age.
Familiesin the "low-quality diet" group were giv-
en the standardseedmixture, and a low-protein nest-
ling mixture (about 7% protein; see Table 1) ad libi-
tum. From the end of incubation on, they received
April1987] Zebra
Finch
Morphology 157

about 5 g of greens once a week, but no egg. Five TABLE 1. Composition(%dry mass)of nestlingmix-
low-diet pairsproducedone broodeach,with 8 male turesusedin experimentaldiets.
and 8 female offspringat 1 yr. All three nestlingfood
mixtures had roughly equal energy content [about Diet quality
3,000kcal/kg or 12.6 x 106J/kg, basedon published Start-
valuesfor componentingredients;metabolizableen- Low dard High
ergy content (Fisher 1972) was not measured]. Rolled oats 34.0
Nine external morphologicalcharacterswere mea- Pablum(babycereal) 24.0
suredon the original parentsbeforethe experiment, Wheatgerm 10.0
on the experimentalbirds as they grew (oncea day Skimmilk powder 20.0 12.0
from hatch until day 14, every 2 days until day 40, Gelatinpowder 10.0 7.6
oncea week until day 98, and at 1 yr of age), and on Vitamin-mineralpowdera 1.0 1.0 1.0
the fully grown offspring later producedby the ex- Alfalfa, dried 1.0 1.0 1.0
perimental birds. Offspring of experimental birds Dry dog food 6.0 20.0
were produced by mating individuals with nonsibs Tryptophan 0.1
Cystine 0.1
from the sametreatment group. Methionine 0.2
The measurementstaken included: mass(MAS) in Wheatgerm 10.0
gramson an electronicbalance;wing chord (WNG) Soyaflour 19.0
in millimeters using a ruler on the flattened right Dried lean beef powder 10.0
wing; retrix length (RET)in millimetersusinga ruler Casein 10.0
from the tip of the pygostyle to the furthest exten- Brewer'syeast 7.0
sion of the tail feathers; tarsuslength (TRS) in mil- Wheat bran 12.0 2.0
Corn starch 68.0
limeters using dial calipersfrom the nuchal notch to
Dextrose 12.0
the furthestextensionof the leg with the foot flexed
at a right angle; toe spread (TOE) in millimeters, aPerveline•, 8 in 1 Pet Products,containsvitamin A palmitate, des-
measuredby pressingthe foot on a ruler and noting iccatedliver, brewer'syeast,wheat germand alfalfa meals,soyflour,
driedmilk,thiaminehydrochloride, riboflavin,pyridoxinehydrochlo-
the distance between the rearmost extension of the
ride, vitamin B•2supplement,niacin,choline dihydrogencitrate,vi-
hind toe and the foremost extension of the middle tamin E supplement,calciumpantothenate,monosodiumglutamate,
toe (excluding claws);bill length at nares (BLN) in tribasiccalciumphosphate,
potassium biphosphate,sodiumchloride,
millimeters from the tip to the anterior rim of the iron orthophosphate,manganese phosphate,magnesiumphosphate,
copperphosphate, zincoxide,potassiumiodide,cobaltphosphate.
nares;bill depth at nares(BDN) in millimeters per-
pendicular to the commissureat the anterior edge of
the nares; lower bill width (LBW) in millimeters at differences between results in all cases. The raw data
the widest expanseof the tomia; body length (BOD) areemphasizedhere,exceptasindicatedin the shape
in millimeters,measuredfrom the tips of the longest and environmentalsensitivityanalyses.Unless in-
retricesto the beak tip of a bird gently flattenedalong dicated otherwise, SAS (SAS Inst. 1982) was used for
a ruler, with the previouslymeasuredRET and BLN statisticalprocedures,with interpretationsbasedon
later being subtracted.All measurements taken dur- Sokal and Rohlf (1981).
ing growth were made by one technician, while I
made all adult measurements.
RESULTS
Growth coefficients were calculated for each char-
acter in each experimental chick from the measure-
Pairs on the low, standard, and high diets
mentsfrom day 1 to 98 in nonlinearregressions based
on a logistic model of growth (Ricklefs 1984). The
had respective averages of 3.2, 2.8, and 2.8
SAS procedure NLIN (SAS Inst. 1982) was used to young surviving to at least one year of age,
estimate K (relative growth rate, as proportion of as- implying that the dietshad no differentialef-
ymptote/day) values, as well as A (asymptoticsize) fect on breeding success.Young producedon
and t• (age at inflection point, or maximum growth the three diets appeared normal in terms of
rate) values. The K values and adult measurements grossbehaviorand appearance,and individu-
for each morphologicalcharacterwere then exam- als from each group went on to breed success-
ined in detail, using two-way ANOVA to examine fully themselves(e.g. Table 4). As is often the
the independenteffectsof sex and diet on growth case in Zebra Finches, there was considerable
ratesand adult size.The effectof diet on morpholog-
variation between pairs within treatment
ical shape was examined using both ratios of char-
acters (Mosimann and James 1979) and principal
groups in the extent to which they used the
componentanalyses(PCA). All variableswere eval- nestlingmixturesfor feeding their young. The
uated for normality before statisticalanalysis. Al- high-proteinmixture was lesspreferredby the
thoughvirtually all analyseswere carriedout on both parentsthan the standardmixture. Somepar-
log-transformedand raw data,therewere only trivial ents given the low-protein nestling mixture
158 PETER
J.BOAG [Auk,Vol. 104

Adults GLM) on the growth coefficients(Table 2)

High
prat
ein showed a consistent,highly significanteffect
of diet on the mean growth rates of all char-
E•e- actersexceptRET. There was essentiallyno sex-
E•o. ual differencein growth rates,and no signifi-
.E cant diet x sex interactions. A factorial
multivariate analysis of variance (MANOVA;

i•[i•/t
I] Low
protein
(x.95
•C.L.)
Age in days
Fig. 1. Growth and adult size of TRS in Zebra
Finchesraisedon low (O) vs. high (0) quality nest-
ling diets.Standard-dietvalueswere intermediateto
low and high values, and are omitted for clarity.
Points are means -+ 95% confidence
are growth ratesbasedon a logisticmodelof growth.
usedit to a greaterextentthan did parentspro-
vided with the standard mix.
choice tests with nonexperimentalbreeding
birds confirmed that adults ranked the palat-
ability of the mixtures as low > standard >
high. The morphologicaleffectsreported be-
low probably derive chiefly from the extra
boiled egg ration in the high diet, vs. heavy
useof the low-proteinmix by parentsfeeding
young in the low-diet group.
Diet and growthrate.--The diet composition
had a cleareffecton nestlinggrowth rates,with
the high diet causingyoung to grow signifi-
cantly fasterthan young rearedon the low diet
(Fig. 1, Table2). Two-way ANOVAs (SASPROC
Norusis 1985) showed no interaction between
diet and sex,no significantsexualdifferencein
growth rates (F = 1.10, df = 10,21,P = 0.41),
and a highly significanteffectof diet (F = 4.96,
df = 20,42,P < 0.001). As an approximatemea-
sure of the effectof diet on within-group vari-
ation in growth rates, ! used a modified Fmax
test (Sokal and Rohlf 1981) on the ratio of the
squared coefficientsof variation (CV2). Varia-
tion amonglow-diet youngwasconsistentlyup
limits. K values
to twice that seenamonghigh-dietyoung,with
the ratio significant in 5 of the 9 characters(Ta-
ble 2; see also Fig. 2).
Diet and adult size.--The experimental diets
Simultaneous had significant,permanenteffectson adult size.
The effect on TRS, a character for which there
is no adult sexual size dimorphism in this
species,is evident in Fig. 1. Similar resultswere
seen for other characters when the measure-
ments of 1-yr-old experimental birds were
comparedwith two-way ANOVA (Table 3). As
with growth coefficients,none of the diet x
sex interactions were significant. There was
evidenceof sexualsize dimorphismin RETand,
to a lesser extent, in TOE. Of the 9 characters,
6 showeda significanteffect of diet on adult
size when sex was held constant. A factorial
MANOVA showed no diet x sex interaction, a

TABLE 2. Effectsof nestlingdiet on means(2) and coefficientsof variation (CV) of relative growth rates(K)
from fitted logisticcurves.

Diet

Low (n = 16) Standard(n = 13) High (n = 15) Test

CV 2
Character • CV œ CV • CV ratio a Diet b Sex

MAS 0.190 29.8 0.255 23.4 0.299 10.4 ** *** NS

WNG 0.243 9.8 0.243 6.3 0.265 4.9 * ** NS
RET 0.307 8.6 0.306 8.5 0.321 6.1 NS NS NS
TRS 0.276 15.0 0.316 12.3 0.370 8.1 NS *** NS
TOE 0.281 18.6 0.336 15.6 0.373 12.4 NS *** NS
BLN 0.124 18.7 0.150 7.1 0.148 9.6 * *** NS
BDN 0.103 16.5 0.120 9.9 0.128 8.3 * *** NS
LBW 0.112 18.2 0.142 12.6 0.139 20.3 NS *** *
BaD 0.212 22.4 0.246 14.2 0.276 9.4 ** *** NS

F testfor largerof Low and High diet CV2 dividedby smaller,with significance
indicatedas *(P < 0.05),**(P < 0.01),and ***(P < 0.001).
Two-way ANOVAs with no significantinteractionterms.Significanceof diet and sexmain effectsindicatedas above.
April 1987] ZebraFinchMorphology 159

TAI•LE3. Effectsof nestling diet on adult size.

Diet

Low (n = 16) Standard (n = 11) High (n = 14) Testb

Character a • CV œ CV œ CV CV 2 Diet Sex

MAS 12.04 12.0 12.40 8.6 13.24 7.3 NS * NS

WNG 55.96 2.5 56.25 2.0 57.12 1.2 * * NS
RET 33.53 3.8 33.53 2.8 34.16 2.4 NS NS ***
TRS 13.19 6.2 13.76 2.4 14.08 3.2 NS ** NS
TOE 18.69 5.9 19.15 4.9 19.96 4.6 NS ** *
BLN 7.85 6.2 7.71 2.0 7.90 2.9 ** NS NS
BDN 7.11 4.9 7.12 3.7 7.43 2.8 NS * NS
LBW 6.71 4.5 6.75 3.1 6.91 4.3 NS NS NS
BOD 65.24 3.8 65.84 2.9 68.81 2.7 NS *** NS

MAS measuredin grams, other charactersin millimeters.

See Table 2 for explanation of tests.

highly significant effect of sex (F = 5.96, df = moved, such componentsdefine ratios of pos-
9,27, P < 0.001),and a significanteffect of diet itively to negatively weighted characters,i.e.
(F = 2.01, df = 18,54, P = 0.02). Relative vari- shape.
ability of the measurementswas comparedin As expected, the PCA on the growth data
high- and low-diet birds as before; again, low- showeda majorPC I dominatedby the overall
diet birds were more variable, though the in- trend for all characters to increase in size dur-
creasewas significantin only 2 of 9 characters ing growth (Table 5). The two-way ANOVA on
(Table 3). PC I scoresgave resultssimilar to the univar-
The morphological differences observed late growth rates;no diet x sex interaction was
among the three groups of experimental birds evident, no sexual difference in PC I (F = 0.05,
at adulthood were not observedin any of the df = 1,30,P = 0.82)wasseen,but a strongeffect
nine charactersin either their parents or their of diet on the growth rate PC I scores(F =
own offspring (typical resultsshown in Table 12.61,df = 2,30, P < 0.0001)existed.PC II rep-
4 for tarsus). resenteda ratio betweenfast-growingleg mea-
Diet and shape:principalcomponents.--Iexam- surementsand slower-growing beak measure-
ined the effect of the experimental diets on ments, or measurements such as RET that were
shapein two ways. First I extractedprincipal not affected by the experimental diets (Table
components from both the correlation matrix 5). The two-way ANOVA on PC II showed no
of growth coefficientsand the correlation ma- diet x sexinteraction,a nonsignificantsexef-
trix of adult measurements. In most sets of avi- fect (F = 3.10, df = 1,30, P = 0.09), and a non-
an morphometricdata,PC I is highly positively significant effect of diet (F = 0.83, df = 2,30,
correlated with all the univariate variables, and P = 0.44). Only PC III showeda significantdiet
canbe interpretedas"size"(Lemen1983,Boag effect (F = 4.22, df = 2,30, P = 0.02), with nei-
1984).The secondand subsequentcomponents ther sex nor interaction terms significant.PC
typically show character correlations of mixed III is not included in Table 5 as it explained
signs; with much of the variation in size re- only 7% of the total variancein growth rates.

TABLE4. AdultTRSsize(in mm,œ+ SE)andANOVA for ZebraFinchesraisedon 3 diets,with correspond-

ing datafor their parentsand offspring.The other8 characters
showedsimilarpatterns.
Diet

Low Standard High ANOVA

Parents 14.01+ 0.13 14.23+ 0.12 14.30+ 0.17 F2,2s= 1.16NS
ExperimentMs 13.19+ 0.20 13.76+ 0.10 14.08+ 0.12 Fz•s = 9.32***a
Offspring 13.79+ 0.21 13.80+ 0.10 14.08+ 0.27 F2,23= 0.68NS
• P < 0.001.
160 PETER
J.BOAG [Auk,Vol. 104

TABLE5. PCAs on correlation matricesfor growth K Coefficients PCA

rates (n = 36) and adult measurements (n = 41).
Entries are correlations between PCs and univar-
iate measures
a (seealso Fig. 2).

Adult
Growth rates measurements

Character PC I PC II PC I PC II

MAS 0.75 -0.07 0.64 -0.46

WNG 0.79 0.04 0.71 -0.05
RET 0.71 0.43 0.56 0.39
TRS 0.82 -0.52 0.83 -0.17
TOE 0.83 -0.46 0.76 -0.05
BLN 0.91 0.30 0.61 0.46
BDN 0.95 0.21 0.84 0.15
LBW 0.75 0.42 0.63 0.43
BOD 0.88 -0.30 0.60 -0.64 -2 -1 0 1

Eigenvalue 6.03 1.06 4.46 1.23 PC I

Percentage
variation 67.00 11.80 49.60 13.60 Adult Measurements PCA
' Boldfacecorrelationsare not significant;all othersare significant
at P < 0.01.

It representedpositive correlationswith WNG

and RET, and negative correlationswith the
beak measures.
The PCA on adult measurements showed that
O -1
PC I summarizesoverall size variation among
adults,with strong,positive correlationson all
characters (Table 5). As for most of the adult -2'

univariate characters,PC I showsno significant

sex or interaction terms, but a highly signifi- -3'
cant effect of diet on multivariate "size" (Table -3 -2 -1 0 1 2
6). In this and other analyses conducted on PC I
similar suites of characters for adult Zebra
Finches, PC II consistentlydefines an axis of Fig. 2. Distribution of PCA scores base;:ton
sexualshape dimorphism. It weighs MA$ and growth-rate variables measured for Zebra Finches
BOD negatively, and RET and the beak mea- raised on low (solid line), standard (dashed line), and
sures positively; male Zebra Finches have a high (dottedline) diets(top). Bivariateroearksfor each
group are indicated(+). Bottomgraph showssimilar
smallerbodythan females,but largerbeaksand results, based on a PCA of adult size measurements.
longer rectrices.PC II in adults had no inter-
L = low, S = standard,H = high.
action term, a highly significant effect of sex,
and a significant effect of diet (Table 6). The
means(Table 6) indicate that the high diet ren- ability (Fig. 2). They accountfor overtwo-thirds
dered the morphology of all birds, indepen- of the total phenotypic variation in growth and
dent of sex, more "female-like," while the low adult data.The experimentaldietsprimarily af-
diet tended to make all birds more "male-like," fected overall growth rate and adult size and
with smaller bodies in relation to beak mea- had smaller effects on shape. The low-diet
sures and RET. PC III from adult measurements younghad morevariablegrowthratesand adult
explained 9.8% of the total variation, and sizes(Fig. 2). Much of the variation in pheno-
showed no significant variation among scores typic responseto the experimental diets ac-
due to diet, sex, or interaction terms. tually was due to between-broodvariation (Fig.
Plots of PC I vs. PC II provide a useful sum- 2). Young from the samebrood tended to clus-
mary of the overall effectsof the experimental ter together,with two pairsof breedingadults
diets on size, shape,and interindividual vari- on the low diet producing young with growth
April 1987] ZebraFinchMorphology 161

T^I•I•E6. Effectsof diet on several indicesof adult shape(œ+ SE). All entries have been multiplied by 10
for display purposes.

Diet ANOVAs a

Character Low (n = 16) Standard(n = 14) High (n = 11) Diet Sex

PC I -5.24 + 2.94 -2.22 _+ 1.42 7.73 _+ 1.43 ** NS
PC II 3.65 _+ 2.66 -1.08 _+ 2.69 -3.32 _+ 2.48 * ***
MASTRS b -7.60 _+ 0.05 -7.75 _+ 0.04 -7.75 _+ 0.05 NS NS
WNGTRS 6.28 _+ 0.05 6.11 _+ 0.03 6.08 + 0.04 ** NS
RETTRS 4.06 _+ 0.07 3.87 + 0.04 3.85 _+ 0.04 ** *
TOETRS 1.52 _+ 0.04 1.43 _+ 0.07 1.51 _+ 0.05 NS **
BLNTRS -2.25 _+ 0.05 -2.52 _+ 0.05 -2.51 _+ 0.05 *** NS
BDNTRS -2.68 _+ 0.04 -2.87 _+ 0.06 -2.78 + 0.04 NS NS
LBWTRS -2.93 + 0.07 -3.10 _+ 0.06 -3.09 _+ 0.06 * NS
BODTRS 6.95 + 0.07 6.80 _+ 0.04 6.89 + 0.04 NS NS
BLNBDN 0.43 _+ 0.06 0.35 _+ 0.06 0.27 _+ 0.03 NS NS
BLNLBW 0.68 + 0.06 0.58 _+ 0.04 0.58 + 0.04 NS NS
BDNLBW 0.25 + 0.04 0.23 _+ 0.04 0.32 _+ 0.04 NS NS

Two-way ANOVAs, with no significantinteractionterms.Significanceof diet and sexmain effectsindicatedby *(P < 0.05), **(P < 0.01), and
***(P < 0.001).
MASTRS is an abbreviationfor log(MAS) - log(TRS).

and adult PC I scoresin the standard-to high- Kruskal-Wallis tests led to identical conclu-
diet range. If a single charactersuchasTRSwas sions.
usedto calculateaveragewithin-broodCVs, the Diet and shape:characterratiosin adults.--Mos-
growth rate meanswere 7.12, 4.31, and 4.93 for iraann and James (1979; see also James 1983)
the low-, standard-,and high-diet young, re- suggestedthat explicit definition of size and
spectively,while the correspondingvaluesfor shape variables is preferable to multivariate
adult TRS were 2.80, 2.00, and 1.81 (given the measuresthat may not be readily comparable
small number of broods, none of the ratios of between studies.They recommendedthat dif-
CV2 were significantin either case).This sug- ferences between the logarithms of various
geststhat although much of the variation was charactersand a single size measure(such as
between broods, the residual within-brood WNG or TRS) are appropriateindicesof shape,
variationmay remainhigher amongsibsreared insofar as they representratios on an arithme-
on low comparedwith high diets. The hetero- tic scale, but take into account the fact that
scedasticityamong treatment groups was not morphologicaldata tend to be log-normally
severe enough to compromise the ANOVAs distributed.
describedearlier, as replicateanalyseson log- I chose TRS as a measure of size in Zebra
transformed data as well as nonparametric Finches.TRS does not display sexual size di-

TABLE7. Correctedexperimentalgroupdeviationsfrom grand means,in SD units.L+H is the sumof the

absolutevaluesof the low- and high-dietdeviations,and providesa measureof environmental
sensitivity
(see text).

Growth rates Adult measurements

Character Low Standard High L+H Low Standard High L+H

MAS -0.81 0.38 0.79 1.60 -0.44 -0.13 0.60 1.04
WNG -0.36 -0.20 0.55 0.91 -0.34 -0.13 0.49 0.83
RET -0.23 -0.17 0.38 0.61 -0.11 -0.16 0.25 0.36
TRS -0.69 0.00 0.85 1.54 -0.65 0.17 0.61 1.26
TOE -0.58 0.24 0.58 1.16 -0.45 -0.06 0.56 1.01
BLN -0.66 0.37 0.48 1.14 0.10 -0.34 0.15 0.25
BDN - 0.76 0.18 0.70 1.46 -0.31 - 0.30 0.59 0.90
LBW -0.71 0.58 0.41 1.12 -0.22 -0.12 0.35 0.57
BOD -0.65 -0.05 0.72 1.37 -0.56 -0.31 0.88 1.44
162 PETER
J.BOAG [Auk,Vol. 104

morphism in this species,has been used by the growth data, the mean deviations for low
James(1983) for the samepurpose,and, along and high diets were -0.61 and 0.61, respec-
with BDN, is the charactermosthighly corre- tively,while the comparablemeansfor the adult
lated with PC I for the adult measurement data data were -0.33 and 0.85. This suggeststhat
(Table 5). The resultsof two-way ANOVAs on the two diets elevated and depressedgrowth
the ratios [actually differencesbetween log- ratesby aboutthe sameamount, while the high
transformed values, e.g. the abbreviation diet increased final adult size more than the
MASTRS indicates log(MAS) - log(TRS)] be- low diet diminished it.
tween each character and TRS, as well as some
ratiosbetween the beak measures,are given in DISCUSSION
Table 6. Once again, none of the diet x sex
interactionswere significant,and only 2 of 11 The quality of food availablefor rearing Ze-
ratiosshowedsignificanteffectsof sex.Four of bra Finch nestlings not only affects growth
11 ratios showed significant effectsof diet, all rates,but also has permanent effectson adult
reflecting the fact that TRS is more sensitiveto size.Low-quality diets increasethe variation in
diet than any of the feather or beak measures. growth rate and adult size both among and
None of the beakshapemeasuresgaveany hint within broods. The possibility that environ-
of diet group differences,indicating that subtle mental effectssuchas the protein level in a diet
shape changes among similar, slow-growing might contribute to spatial or temporal mor-
characterswere not taking place. phological differences in birds has been con-
A measureof environmentalsensitivity.--To sidered by only a few workers to date (Smith
compare the effectsof the experimental diets and Zach 1979; James1982, 1983; Boag 1983;
on the nine charactersfor both growth rates Alatalo and Lundberg 1986). To interpret the
and adult measurements, with sexual dimor- results of my experiment, it is important to
phism statistically removed, a standardized consider first if the small differences created
measureof sensitivity to the dietary manipu- here could be of ecologicalsignificance,and,
lation was created. First, all the growth coeffi- second,whether suchdifferencesare likely to
cient and adult measurement data were con- be produced under natural conditions.
verted to Z scores (mean of zero, standard To evaluate whether the morphologicaldif-
deviation of one). Using the SPSS-Xprocedure ferencesproducedhere could be of ecological
ANOVA (Norusis 1985), two-way ANOVAs on importance, consider the difference between
diet and sexwere performed with MCA (mul- high- and low-diet means in Table 3 as a per-
tiple classificationanalysis).Such an analysis centageof the low-diet means.The effectof the
gives the deviation from the grand mean of high diet is to increasethe low-diet adult means
each experimental group, in standard devia- 1-2% for BLN or WNG, or 7-10% for characters
tion units, after adjusting for the other main such as TRS or MAS. Changesin means of less
effects. Thus, in the case of an adult measure than 2% are difficult to detect in nature without
such as TOE, diet explained 10% of the total large samplesizes,and to detecttheir function-
phenotypic variation, with means for low = al impact on activities such as foraging (Schlu-
-0.45, standard= -0.06, and high = 0.56 (sex ter and Grant 1984)would be equally difficult.
held constant), while sex explained 9% with Effectsin the 3-10% range could be detectedin
means for males = 0.25 and females = -0.35 the wild, and are more likely to be of function-
(effect of diet held constant). The measure of al significance.Indeed, many avian studieshave
environmental sensitivity chosenwas the sum assumedthat intraspecificmorphological dif-
of the absolute values of the corrected high- ferencesin this range are important.
and low-diet deviations from the grand mean For instance,many monogamouspassetines
(L+H values in Table 7). displaysexualsize differencesof lessthan 10%.
There was a significant correlation between The significant differencesbetween survivors
the L+H values basedon the growth data and and nonsurvivors in the Bumpus House Spar-
the adult data (Spearman rank correlation, n = row data (O'Donald 1973) and Isla Daphne Ma-
9, rs = 0.73, P < 0.05). Characters such as MAS
jor Geospiza
fortisdata(Boagand Grant 1981)lie
and TRS showed consistently large effects, in the 2-8% range. In examplesof ecotypic
while LBW and RET showed small effects. In variationin avian morphology,Fretwell (1972)
April1987] Zebra
Finch
Morphology 163

noted significant between-habitat variation in under natural conditions.In general, we do not

tarsus length for Savannah Sparrows (Passer- have adequate data to answer this question
culussandwichensis,10%),Song Sparrows(Melo- confidently.A few field studieshave reported
spizamelodia,4%),and Field Sparrows(Spizella seasonalor annual variation in morphology
pusilla,2%).Grant et aL (1976)found significant (Perrins 1979, Smith and Zach 1979, Boag 1983),
between-habitat differences in Geospizafortis and Alatalo and Lundberg (1986) found signif-
morphologyof well under 10%,while morpho- icant differencesin tarsuslength of Pied Fly-
logicaldifferencesof a similarmagnitudehave catcher(Ficedulahypoleuca)offspring reared by
been noted for titmice (Parus spp.) living in primary vs. secondaryfemalesof polygynous
coniferous and hardwood forests (Perrins 1979). males. The differences (about 2%) were attrib-
Authorsstudyingclinal variation in avian mor- uted to differences in paternal feeding rate.
phology (James1970) or making island-main- Alatalo and Lundberg(1986)alsofound signif-
land comparisons(e.g. island subspeciesof the icant effects of breeding density on tarsus
wren Troglodytestroglodytes; see Williamson length, with nests in high-density areas pro-
1981: 139) also commonly discussthe adaptive ducing smaller offspring.Several other long-
significanceof differencesbetween popula- term studiesprobably have similar data, al-
tions of well under 10%. In a recent summary though often birds have not been measuredas
of temporal and geographical morphologial adults, or different workers have been in-
variation in the Fox Sparrow (Passerellailiaca), volved in different areasor years,complicating
Zink (1983) sought evolutionary interpreta- comparisons.
tions of results for which the largest average Despite the paucity of data, there is no rea-
percentagedifferencewas 3.28%,with a range son to believe such effects are rare in nature.
of 0.55-8.47%. Adaptive explanationsfor sig- The experimentalregime I used does not in-
nificant morphological changes of 3-5% in volve a totally unrealisticor pathologicalrange
House Finches (Carpodacus mexicanus)in their of diet qualities. Many estrildid finches rou-
new easternpopulationscomparedwith their tinely raise young on seedalone in the wild
ancestralwestern populations were proposed (Goodwin 1982, Morton and Davies 1983, Zann
by Aldrich (1982).Dhondt et aL (1979) found and Straw 1984), and many of them inhabit sa-
that Great Tits graduallybecame3-7% smaller vannah habitats with concomitant seasonal and
over 14 years.Severalmechanismsfor the shift annual unpredictability in the timing, quanti-
involving natural selectionwere suggested,re- ty, and qualityof resources
during the nesting
volving around the fact that the addition of season.This may result in occasional,oppor-
nest boxeshad doubled or tripled Parusbreed- tunisticinsectfeeding or periodic food limita-
ing densitiesduring this period. tion, during which time nestlingsand fledg-
In citing these studies, I am not suggesting lings starve(Zann and Straw 1984).Granivores
that all such examplesinvolve morphological with faster development times than Zebra
differences attributable to nonadaptive, envi- Finchesrequire a high-protein diet for breed-
ronmental effectssuch as nestling diet quality. ing to be successful.Such speciesoften feed
I merely point out that avian ecologistsregu- nestlingson insects(Jonesand Ward 1976) or
larly seekadaptiveexplanationsfor differences other protein-rich itemssuchasthe specialized
of a magnitude that I have producedby way of use of Spirogyraalgae by Sharp-tailedMunias
nonpathologicalvariation in diet quality (by (Lonchura striata;Avery 1980). Many birds de-
nonpathological I mean diets not producing pendent on insects for rearing young show
nestlingdeathor infertile adults).In somecases, considerable seasonal or annual variation in
e.g. the trend seenby Dhondt et aL (1979) in nestling growth rates, fledging success,and
Parussize over time, I see parallels with the brood-reductionbehavior (Bryant 1975, 1978;
breeding-density-associated annual variation in Perrins 1976; Howe 1977; Ricklefs and Peters
offspring size noted by Boag(1983) for Geospiza 1979),implying that the potentialexistsfor diet-
fortis. induced morphologicalchange. Dietary pro-
The second question raised above was tein can have major effectson both growth and
whether variation in diet or other aspectsof adult size of captive ducks (Johnson 1971) and
the nestling environment sufficientto produce domestic fowl (Fisher and Griminger 1963).
significantmorphologicalchangesare common Other environmental effects have also been in-
164 PETER
J. BOAC [Auk,Vol. 104

vestigated.For example, pullets raised at 6øC tween-broodmorphologicaldifferencesthat are

during 3-6 monthsof age were shorter,gained due to environmental effects. Some effect of diet
more weight, and had smaller tarsi and tails on variation within Zebra Finch broods re-
than former flock mateskept at about 23øC(A1- mained after accounting for between-brood
lee and Lutherman 1940). Similar studies re- variation, indicating that processes
suchas sib-
ported the impact of environmental effectson ling competitionmightaffectintraspecific mor-
morphology in mammals (Widdowson and phologicalvariation.Environmentaleffectsthat
McCance 1960, Lister and McCance 1965, Wil- alter the distribution of variance within and
liams and Hughes 1975, Barnett and Dickson between broods can have important conse-
1984), including humans (Hulse 1968). In one quences for heritability analyses (Falconer
of the best longitudinal studiesof associations 1981).Althoughsucheffectsare citedregularly
between intraspecificmorphologicalvariation in criticismsof heritability studiesunder field
and fitness in mammals, Clutton-Brock et al. conditions,almost no data exist to support their
(1982)found that diet or density-relatedmater- existence or nonexistence. Van Noordwijk
nal effectscan have profound effectson body (1984)suggested that environmentaleffectsre-
size and the lifetime reproductive successof suitingfrom naturallyoccurringpoorbreeding
individuals or entire cohortsof red deer (Cervus seasonsor from manipulation of brood size can
elaphus). at least temporarily increase morphological
Among the Zebra Finches used in this ex- variation among sibs,and hence decreaseher-
periment, there was considerablevariation in itability estimates.
the responseof different breeding pairs to the The Zebra Finch data suggestthat the pri-
diet treatments;one pair wasableto raiseyoung mary effectof variationin diet quality is on
to "high-diet size" on the low diet. Part of this overall growth and adult size. There was not
variation may be an artifact of the multicom- enough differential impact on charactersto
ponent, qualitative diets used, which made it produce large changesin shape. The small
difficult to control the amountsof protein, en- shapechangesobservedwere due primarily to
ergy, or other dietary componentsconsumed differences between characters in their rate and
by each young bird. Zebra Finchescan raise timing of growth. Measurements
suchas MAS
young on seedalone, so it is unlikely any birds or TRS increasequickly and reach asymptotes
sufferedfrom an energyshortfall.Similarly,all at an early age; they are affectedmostby vari-
nestling and egg mixturescontaineda vitamin- ation in diet. Beakcharactersgrow more slow-
mineral supplement,making specificshortages ly, for a prolongedperiod.Beaksare lesssus-
in these componentsunlikely. The most ob- ceptible to dietary influencesthat have their
vious difference between the diets was in crude majoreffectwhile youngaredependenton their
proteinavailability;however,in the absenceof parents.Charactersinvolving feather growth
detailed data on assimilation efficiencies, ami- also are affectedless seriously,despite rather
no acid balance,or micronutrient availability high growth rates. This reflectsthe high de-
(Fisher 1972), it would be misleading to con- velopmentalpriority of passerineplumagede-
clude that the morphologicaleffectsobserved velopment, which tends to proceedaccording
here are solely the result of dietary protein to a fixed schedule even if a nestling is starv-
level. Some of these difficulties could be avoid- ing.
ed if a single, synthetic food ration had been It remains to be seen if environmental effects
used(e.g.Murphy and King 1982).It is possible inducedby variationin the quality or quantity
to switch Zebra Finches to a synthetic diet, but of nestlingdiets, or other aspectsof the nest
this can causeadult mortality and poor breed- environment, regularly influence natural pat-
ing success, at leastinitially (T. Scheuhammer terns of intraspecificmorphologicalvariation
pers. comm.). The complications described in birds. Considerablescopeexists for long-
aboveare importantin designingfuture exper- term, longitudinal studiesof the environmen-
iments, but do not alter my principal finding tal correlates
of morphological variation,aswell
that diet quality can affectadult size as well as as more natural experiments of the type I have
growth rates in Zebra Finches. conductedhere (e.g. Berthold 1976, 1977;Smith
Under natural conditions, differences in the and Arcese 1987). The primary data on varia-
quality of parental care might produce be- tion in avian growth rateshave not been sup-
April 1987] ZebraFinchMorphology 165

portedby studiesthat followed youngto adult-
hood to check for permanent effects on
morphology. Hence, we cannot discriminate
BOAG,P. T. 1983. The heritability of external mor-
which avian charactersdisplay compensatory,
or "target-seeking,"
growth (Atchley1984)once
the period of parental dependenceis past,and
which do not. Judicious choice of characters
may well minimize the likelihood of signifi-
cant environmental effects in studies
morphometrics.Until additional data are avail-
1977. Ober die kiinstliche Aufzucht Nest-
junger Amseln (Turdusmerula)reit Beeren des
Efeus (Hederahelix).Vogelwarte 29: 110-113.
phology in Darwin's ground finches (Geospiza)
on Isla Daphne Major, Galapagos.Evolution 37:
877-894.
1984. Growth and allometry of external
morphologyin Darwin'sfinches(Geospiza)
on Isla
of avian Daphne Major, Galapagos.J. Zool. London 204:
413-441.

able, however, the possibility of environmen- , & P. R. GRANT. 1981. Intense natural selec-
tal effects should not be dismissed automati- tion in a population of Darwin's finches (Geo-
cally in favor of selective explanations for spizinae)in the Galapagos.Science214: 82-85.
ß& A. J. VAN NOORDWIJK.1987. The quanti-
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ACKNOWLEDGMENTS Eds.). London, Academic Press.
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I thank Radiq Tallim, Jody Morgan, and my un- Martins Delichon urbica in relation to aerial insect
dergraduateassistantsfor their help. L. M. Ratcliffe, abundance. Ibis 117: 180-215.
J. N.M. Smith, and the refereeshelped improve the ß 1978. Environmental influences on growth
manuscript.My Zebra Finch researchhas been sup- and survival of nestling House Martins Delichon
ported by researchgrants from Trent and Queen's urbica. Ibis 120: 271-283.
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G. A., JR. 1979. Bodyweightsof birds:a re-
the Natural Scienceand EngineeringResearchCoun- view. Condor 81: 193-202.
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