Biu/o@cn(Jou~nal oJthLinnmi .Soriep (2000). 70: 27-36. b ' i t h I f i p r r
doi: 10.1006/bijl. 1999.0362, available online at http://ww.idealibrar)i.com on
1DE b1@
Fluctuating asymmetry in antlers of fallow deer
(Duma duma): the relative roles of environmental
stress and sexual selection
R. J. PUTMAN" AND R/I. S. SULLIVAN
Behavioural and Environmental Biology Research Group, the Manchester hletropolitan
Universip, John Dalton Building, Chester Street, Manchester M I 5 G D
J. LANGBEIN
Dzvision of Biodiversip and Ecolog, Universip of Southampton School of Biological Sciences,
Boldrewood Building, Bassett Crescent East, Southampton SO1 6 7PX
Recezved 14 Octobn 1998; acrpptcdfor publication 25 March 1999
Antler lengths were recorded of a total of 250 male fallow deer (Duma duma (L)). Animals
sampled were from marked populations where the majority of individuals w-ere of known
age. Asymmetry in antler length was normally distributed with a mean not significantly
different from zero? confirming that differences in length between the antler pair constitute
a true fluctuating asymmetry (FA). We found no clear relationship between the degree of
asymmetry in antler length of an individual male and either population density or actual
body mass. We did, however, detect a significant relationship between asymmetry and
deviation from maximum cohort bodyweight. If deviation from maximum weight within a
cohort may be considered some index of competitive success or increasing environmental
stress, this may suggest that asymmetry in antler length relates in some way to developmental
stress suffered by the individual concerned. The degree of asymmetry recorded in antler
length also showed a significant decline with animal age, with antlers of animals of 2 years
or older showing significantly greater symmetry. This is consistent with a hypothesis that
despite continued competition for resources, there is a changing balance of selection pressure
as animals reach maturity, with increased pressure from sexual selection requiring males to
produce significantly more symmetrical ornaments.
0 2000 The Linnean Society of London
ADDITIOXAL KEY WORDS:-fallow
deer
~
antlers
-
fluctuating asymmetry.
CONlENTS
Introduction . . . . . . . . . . . . .
Methods . . . . . . . . . . . . . .
Results
. . . . . . . . . . . . . .
Antler asymmetry in relation to development
Changes in antler asymmetry with age
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'stress'
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* Corresponding author. E-mail: [email protected]
002~-4066/00/050027+10 $35.00/0
27
0 2000 The Linnean Society of London
R. J. I’CT\I.\N 67 -11.
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;i\x.n ttiv h r s i s that the ultimate cxpt-rssion of ail)- physical characteristic is ;I
tiiiiction h t l i ot’ gerictic potential and of’ eiiviroiinictital factors aiTccting the dc\ ~ I o ~ ~ i i i cof
i i tthat character. recent studies of fluctuating asy~niiictr!~il.’;Zj I i a \ - c
1; ) c . r i ~ c do n the, fbllo\ving concepts:
I
11
...
Ill
ii\,
that hilatcrally or raclially synirnetrical structures are rarel!. clt.\.elopcd iii
:il)solutc s!-riirneti~ but usually displa)- some sniall residual ;i>!mmetr!. i r.g.
I,ud\z.ig. 19311; van Valcn. 1962; hloller & Swaddle 1997):
that the relative clcgrer of asymmrtr!- ol~scn.edin different indi\~iclual.;\i-itliiii
pqiiiIatiotisTor twtiveeti populations of tlir sarnr organism, nia)- tliiis reflrct
tliffcrcnces in the degree of dc\~lopmental‘strrss’ sufTcrecl tiy cliff erent incli\.idrials/l,opulatioiis [Leary & Xlleiidorf, 1989; Parsons, 1990; Clarke. 1 ()!
Iiozlov (’t 01.. 1996; A~11)llc.r& S\\raddle. 1997);
thal \\.ithin populations. those incli\.iduals best a l ~ l cto rsprrss true .;!mimeti-!.
c I ( y i i r tlic cle\-eloprncntal stress, are likel). to lie the fittest and that t h l i h
approsinlate sj-mmctry may lie used in mate selection as an indicator of fitnc.s:s
(Alollcr, 1 S92; Moller Sr Poniiankoivski, 1993a; \\‘atson 8: Thornhill, 1 cW:
C:larkc, 1W5):
that if symmetry is taken as an indicator of mate qukalit),. sexual selectioii
sliould strotigly fa\.oiir symmetry in particular in sexual display c l i a r a c t c ~
ihloller & Pomiankoiz,ski, 1993h; hloller & S \ \ d d l e , 1997).
If the dcgrcc of fluctuating as);nimctry is to he taken in this \va). as an indci o f
quality. it is lio\ve\,er crucial to rstal~lisli: (1j that the initial t e n d e t i c ~t~o
as! tiinietr)’ i h indeed a reflection of cn\~ironmentalstressors, and (2j t h a t \\.liichc\.c.r
scx rsrrcises choice of mate does indeed sclect those coming closest to true symtncrr!;or inclirectly that in rcsponst. to this, selection is serti to act to reduce asyrnmtw!
in rc]iroducti\~eadults). In such contest the antlers gro\vii and shed througliout liti.
1)) m;dr drcAr pro\kle a11 csccptiorial system in \\.hicli t o rsplore these c ~ i t c s t i o ~ i s ~ s i t i ~ ~ e
,iiirlcrs are tleciduous structures, gro~vnagain in inore complex form each ).ear--;tncl
rlic.) .;lion- distinct arid easily measurable asymmetry ((’.g. Solbcrg Kr Sacthei., 1 PO?;:
ll:il!.oii R Hcaly, 1994; hfarkusson 8r Folstad, 1997).
llcilc iillo\i. deer (Domn dmrr (L) gro\v their first set of antlrrs as ycarliiig\,
Yc.;ii-lings ‘arc t h u s of particular interest in assessment of the role of rii\.ironnieiit~iI
stwsmrs it1 de\~elopmentalasymmetry-hut not siitipl). liecause this is the first antler
gr(m d i . In m a n y other respccts as \veil, ).carling males prove as a class particulat-l!wtisiti\.e t o prc\.ailing envirotimental conditions (dcnsit)-,food qualit!-. food cjuantityr.
111 c.pisode.: of lira\-). ovcnvinter mortality. male yearlings suffer a disprop rtion;itcl!
high inexsure of mortality in comparison t u other age-/sex- classes i n both r t d d t ~ r .
(;(,/i’/d\ t l ~ / ) / / / iatid
\.
fallow derr, Dan20 L’O~/O (Glutton Brock & “Ubo11. 1982: Cllitton
131x)(.k.Giiiniirss 8: Albon, 1982; Putman R Langbein. 1992a). Even hcforc thest. arc.
cipr(xwd in increased mortality. changes in food qualit)- or density are imtiiediaic1~rctlcctcd i t 1 c,lianges in lmd\ivcight of tliesc yearling males, \ w l l M u r e ~ u c hcfh.ts
apparrmt in any other class of individuals (see fix example Langt)rin. 1W i :
iiiiite
Langbein 8: Putman, 1992; Putman & Langbein, 1992a). Thus in effect these
);earling males are especially sensitive to environmental 'condition-or
stressors
influencing the population as a whole. Antler growth in yearling males thus prolides
an excellent system in Izliich to explore the relationship 1)etLveen environmental
stress and expressed asvmnietry. Our hypothesis here (H)-pothesis 1) is that the
degree of asymmetry of different populations or individuals will reflect some measure
of environmental stress such as population density, food quality or condition as
reflected in absolute body mass.
A study of antler growth however also permits exploration of our second question:
the cstent to which sexual selection stri\;es to override developmental stress, and
promote greater symmetry among males of reproductive age. hlale deer cast and
regrow their antlers every year, so each new set of antlers reflects the environmcntal
conditions over the period of deposition. HoLvever, in addition, as animals get older,
the balance of differing environmental pressures shifts as sexual selection starts to
exert a stronger influence.
In the majority of more social sprcies of deer (red and fallow, as well as sika deer
Cemus ~z$pon) mating systems are strongly polygynous (Putman, 1988) \\;ith a very
pronounced skew in mating success apparent amongst mature males (Clutton-Brock
et a/., 1982; Peniberton et a/.> 1992; Langbein, 1991; Thirgood, 1991; Thirgood,
Langbein & Putman, 1999). Female choice of mate is affected at least in part by
body weight and antler size (Clutton-Brock et a/., 1982, 1989; Clutton-Brock &
AlcComb, 1993; Apollonio, Festa-Hianchet & Rlari, 1989). Selection thus fa\.ours
among mature males the niost rapid development of the largest set of antlers they
can achieve; at the same time, strong competition between males enhances the skelv
in dominance and thus access to resources. 'Stress' might then be exaggerated both
through greatcr need (more rcsourccs required to grow bigger antlers) and enhanced
competition; we might thus expect a change in asymmetry in relation to age, or in
reproductive vs immature individuals.
Indeed, if emironmental stress is the dominant factor in determining the degree
of asymmetry expressed, one might expect a U-shaped cur1.e of asymmetry \:ersus
age, xvith yearling males displaying high levels of asymmetry, adult but non-breeding
males with better resource access showing lower le\.els of asymmetry, and with
asymmetry increasing again in the highly competitive reproductive age class (Hypothesis 2A). However, if the degree of synimetr) which may be achieved is a clear
indicator of fitness, and if females select males in part also in relation to antler
symmetIT (e.g. Solberg & Saether, 1993) then sexual selection might be expected
to maintain good symmetry even amongst reproductive males, despite the increased
competition during del~elopment(Hypothesis 2B).
In this paper Lve examine the expressed asymmetry in antler length of fallow deer
from nine different populations, using data on yearling and adult males to test these
two hypotheses. We show first that differences in antler length on left and right
sides are a true fluctuating asymmetry (Sullivan, Robertson & Aebischer, 1993;
Palmer, 1994; Swaddle, Witter & Cuthill, 1994) and not a functional and directional
asymmetry as suggested by tU\:arez (1995). We then ask: is asymmetry in antler
length related in any simple way to measured environmental stressors at the time
of growth? Finally we compare le\.els of asymmetry obser\:ed in antler length of
males of dfierent age or reproductive status, to ask the supplementary question:
Does antler asymmetry increase or decrease with age and reproductive competition?
Data on overall antler length were collected for a total of 250 individual fallow
deer (125 yearlings; 19 known 2 year olds; 12 known 3 year olds; 94 adults of 4
)ear\ or greater) from 9 enclosed @ark) populations, during routine live-capture
operations in the course of other work (see Langbein, 1991; Langbein & Putman,
1992; Putman & Langbein, 1992a, b). Antler lengths only were recorded and no
attempt u as made to investigate potential asymmetry in antler shape, palm width
or number of spellers; while such variation in overall shape has been considered by
other authors (e.g. Solberg & Saether, 1993; Malyon & Healy, 1994) most relationships detected appear to be primarily with simple length. Further, yearling
males produce simple, unbranched antlers with no complexity of structure; since
dnalyses reported here depend on comparisons between yearling and older males,
0111) measurements of simple length provide the necessary consistency between agecl'mes.
Animals were caught in longnets (Cockburn, 1976; Smith, 1980) or fixed handling
units (see Langbein, 1991 for details) and catches were undertaken over winter
(December-February) from 1985 to 1992. All animals captured were weighed on
an electronic weighing platform, aged and marked for future identification with
coloured numbered eartags. Animals were aged on capture as fawns of the year
(thus between 6-8 months of age), yearlings or adults. Animals first caught as mature
individuals were merely classed as adults of unknown age; however since capture
operations continued over a number of years, a number of animals first caught and
marked as fawns or yearlings could, when recaptured subsequently as adults, be
more precisely aged to 2, 3 or 4 years.
Antler lengths were recorded using extending vernier callipers to measure the
straight line euclidean distance between the base of the pedicle and the furthest antler
tip. Animals with broken antlers were excluded from any analysis. Measurements were
recorded to the nearest mm. Repeat measures for confirmation were not undertaken
(Swaddle et al., 1994) but the scale of difference in length between right and left
antlers (with a maximum recorded difference as much as 292 mm) greatly exceeded
estimated measurement error (estimated at <0.5 mm).
In all populations, total population size and age-structure were known from
capture and marking operations and direct counts (Langbein, 1991). Since study
populations were emparked within known areas, densities could also be calculated.
In practicc two density figures were calculated as:
(1) absolute densib (simple numbers per unit area, or, where deer shared the park
land with other grazing stock (other species of deer, or sheep) as livestock units
per total area; see Putman & Langbein, 1990, 1992a); and
(3) e$ectir*e dens^& (livestock units calculated per actual area of available grazing
<ground).
From capture operations, individual weights of marked deer were available, and
average weights could be calculated each year for each age-/sex- class within the
population as a whole. Following Putman & Langbein (1990, 1992a) we used mean
yearling male body weight as a further index of population condition overall.
M.'e calculated a measure of:
(1 j
agmrnetty as the difference between the lengths of the two antlers, divided by
AGE AND ASYRlhlETRY IN E’.\LLO\V DEER ANTLERS
31
the length of the longer. This measure of asymmetry was selected amongst
the range of possible alternatives after exploration of relationships between
asymmetry and 01 era11 antler length, and following recommendations of Moller
and Swaddle (1 997);
(2) deviatzori from rnuwnum agr-class wight as the difference between an individual’s
own weight and the weight of the heaviest animal in the same cohort in the
same population, divided by that maximum weight (all calculations undertaken
separately for yearlings, known 2, 3 and 1 year olds, and all adults);
(3) devzationfrom mean age-rluu te’eight, as above, but the (positive or negative value)
of the difference between an individual’s weight and the mean weight of that
age-class in that population, again as a proportion of that same mean weight.
Since neither density nor effective density of the study populations changed
significantly over the period of study (Langbein, 1991), in all analyses at the
population level (whether within or between populations) antler measurements taken
in different years in any one population were subsequently pooled for analysis.
Because capture operations were conducted in the same parks over a period of
years, a number of individual animals in each population were recaptured on
successive occasions. In order to eliminate risk of non-independence of data, only
the measurements taken the first time each of these individuals was captured as an
adult are included in population level analyses. Repeated measures of the same
individuals in successi\ e years were however included in a separate analysis of
changes in antler asymmetq within an individual over time. Eleven males were
measured in two successive winters; four males were caught and measured in three
successive years.
RESULTS
A clear asymmetry in antler length was apparent in all age-classes of animals
measured and from all populations. In contrast to AIvarez (1995) we found no
consistent ‘bias’of left-handedness or right-handedness, with no significant difference
in the number of animals with left antlers longer than the right or right longer than
left. In our samples 107 individuals had right antlers longer than the left, 21
individuals had antlers of equal length, 119 individuals had right antlers shorter
than left. Further, the antlers of known, marked individuals, as commonly alternated
in successive years in which was the longer side as remained consistent with one
side consistently longer throughout.
When attributed with sign, distribution within the population of the calculated
measure of asymmetry was clearly normal around a mean value not significantly
different from zero (x = 0.006; SD = 0.14; .N= 250; z = 0.01, P= 0.7 1).
antler asymmety in relation to developmental ‘stress’
In Analysis of Variance between populations, there were no significant relationships
in the degree of asymnirtry among yearlings in relation to either measure of density
used or the ‘index of condition’ provided by mean yearling weight (ANOVA on
antler asymmetry by density class; by mean cohort weight: N= 125, F= 1.08, P=
g 0.09
@,
'
0.08
0.07
0.06
1
0.05 ;
I
004
I
T
7
1
-I
L
3
4+
I
2
1
Age class (years)
O . : M ) .Nor a s any relationship detected bct\veen an individual yearling's asymmetry
a i i t l its o\\.ti Iiodyt\'eight or the deviation (if its Lveight from the mean or masimutii
\\rights rccordecl for its cohort (simplc rcgrcssion; ,V= 125, F= 2.56.P= 0.1 12; F =
1 .!)~5. P= 0.1(i5. respec'ti\d!,).
Tlic, .;aim analyses were also uiidertakrn with older animals. Once axain, there
\ \ a s n(1 c\.idcncc for a relationship between an individual's asymmetry and its actual
l)od!iveiglit, or the weight deviation from the mean or maximum recortled for its
ag:c (,lassiii the particular (park) population from which it was drawn: no rrlationship
\\;is detectcd for any age-class, whether for animals of 2 years, 3 ycars. or in the
largcst sample size of animals of 4 years.
Thcrc ivas. liowwrer, a significant p0sitiL.e relationship between asymmetry and
tlr\-iation tiom maximum bodyweight for the age-class when all males wcrc c-onsidcrcd
toget her (pooled analysis for all age classes, yearlings as well as all adult niaics: ,\.=
228: F= 4.65;P= 0.03).
+
'The degree of antler asymmetry recorded showed a significant change \I ith ariinial
age (Fig. 1 ; animals from all populations p o l e d ; Kruskal-Wallis test on asymmetry
t)!- agecl:taa;,\'= 250,1age-classes, H = 16.9 1. RO.00 1). Yearlings sholvrci far greater
a 5 y i m e t r ~than
~ other age-classes; no significant differences in degree of asymmetq
\I-(%IYapparriit lietween animals of 2, 3) 1
years (.N= 122, P= 0.80). If anal
rc.pc;ttrtI, hut restricted to animals of accur-atelydetermined age (those kno\vn to he
1. 2. 3 01'L )'ears of age) the same trend remains, with yearlings ha\-ing far more
jxoiioLiiiccc1 asymmetr)' than animals of 2,3 and 3 years, but sample size is much
sinallcr atid the result is no longer significant (,V= 157; P= 0.13).
Xlatclied pairs tests (\Vilcoxon hhtched Pairs, signed ranks) also failed to show a
+
significant directional trend of changes in asyninietry in antlers of individual males
measured in two (h‘=11) or 3 (“Y=4) successive years (total year-pairs = 19: T =
80; for P=0.05 TI.,,,
46).
DISCCSSION
Asymmetry in recorded antler length of fallow bucks was normally distributed
with a mean not significantly different from zero; antlers of individual males measured
in successive years showed a reversal of bias as often as a tendency for one side to
be longer than the other. These results confirm that differences in length between
the antler pair constitute a true fluctuating asymmetry and show no evidence of
directional asymmetry as suggested by Alvarez (1995).
Our analyses in this paper address two main questions:
(i) Whether asymmetry in antler length is related in any simple way to environmental stressors at the time of growth (density, food quality etc.), providing
evidence that the relative degree of fluctuating asymmetry observed in different
individuals within populations, or between populations of the same organism,
may reflect the degree of developmental ‘stress’ suffered by those individuals/
populations.
(ii) Whether antler asymmetiy increases or decreases with age and reproductive
competition and whether we have evidence to suggest that sexual selection acts
to impose greater symmetry on reproductive animals, suggesting that degrees
of asymmetry do indeed offer an accurate reflection of fitness.
We found no clear relationship between the degree of asymmetry in antler length
and actual body mass (controlling for age). We did, however, detect a significant
positive relationship between asymmetry and deviation of an individual’s weight
from the maximum for its cohort when all age-classes were pooled in analysis. This
same trend was apparent when analysis was restricted to yearlings only, although
with this more restricted sample it did not reach significance (P=0.165, with .V=
125). If deviation from the potential maximum weight of its age class may be
considered a measure of decreasing individual competitive ability, or increasing
environmental stress, this relationship might suggest that asymmetry in antler length
does indeed relate in some way to developmental stress suffered b) the individual
concerned.
For moose A c e s alcts Solberg & Saether (1 993) also report a negative relationship
between antler asymmetry and body inass when controlling for absolute antler size
(thus in effect exploring the relationship between antler asymmetry and body mass
in animals of equivalent antler size). Thus relatively larger bulls in relation to their
antler size produced more symmetrical antlers compared with lighter bulls of the
same effective cohort, a result thus entirely consistent with that presented here for
fallow deer.
‘The degree of antler asymmetry recorded in fallow deer also showed a significant
change with animal age. Yearlings show far greater asymmetry in antler length than
other age-classes, supporting the concept that yearling males are more sensitive to
any environmental stresses than any other class (Putman & Langbein 1990, 1992a,
b). Although body growth and somatic skeletal development is not completed in
male fallow deer until approximately 3 years of age (Langbein, 1991), growth rates
decline significantly after the first year. Demand for resources for skeletal
$1
I<. J. p(..i'.u.\s
F r .I[.
(inc~lopnicnt,particularly minerals, is t h u s especially high for yearliiig niales \vho
prolxhly crperieiicc s e ~ ~ restriction
rc
of thosc rcsources spccificall! requit-td l i ~ r
Ill) I l Y yu\\dl.
. \ tlecliiir in as)ninictry with age is also consistent with t h c sccoiid h>ptliesis t o
tc.tcci iri this paper: that if femalc choice is itifluenccd by aiitler syiinietr)-. scsual
v l c i . t i o i i iiia). ~tk111and'niorc .;ymmctrical antlcrs in iiiorc n i a t u t ~niales. clcspitr
s o i l i t ' c.ontiriued coinpetitioii for- resources espcrienced in devc%lopmeiitof each lie\\.
\ (,;it.'>
gIo\\di. Ho\vc\.er, fa ll o~itnalcs
~
rarely liecome scsuall!- acti\,r ;ind successfiil
i t i 1 onipt~tingfor niatcs ~intilthey arc fiilly mature at 4 or 3 yean of. agc. The
.;igiiificaiit drop in asymruetry is obscr\,ed between yearlings and aiiinial~of two
\ ( ' a r a nf q t . , \\it11 no significant difference in asymmetry rccordcd Iiet\\.ccn 111o( ' i i I' old.; ant1 older bucks. ?'his \ v o u l d perhaps suggest ii lesser rolc of .;c.sual srlectioii
a i i t l a r p c tliat thc decline in asytnmrti-y ma)- he priiiiari1J- due to ;i reduction iii
lopmcnt once anim;ils h a w p i
i ~ t i \ii.oiiiiiciita1 styes\ o \ ~ the
r pcriotl of antler d
tIi(, (,ritic,;il>-earlingphase.
. \ i i inc,i.c.ascin syinniet? of antler (palm) length in fallou- deer kvas also f o ~ i n dt o
c I )rix>Iatc\\.it11 Imth shouldei. lieiglit (thus age) and beha\~iouraltiieasui-es o1'dorniti;rriic~t~
I)!. hlalyoii 8r Heal) (1 994) w h u , while \\mking o n l ~ivith
.
a small aaniplc of' eight
I )~ic.ksfound that niore dominant males Lvithin their study population had t)oth
liirgt~,and 1nc)rc symmctrical antlers ( a s ~~cprrsentcd
hy the relati\.e lengths of' left
a t i d right pilnisj. A similar ne pti \-e relationship hetiyeen ()\,era11 antler size and
criiiiplrxi~y\thus age) and asyinmctry has been reported for rcindccr t)!. 31arl;ii.;son
tk Folstad j 1!397). By- contrast S o l l ~ r g& Saether (1993) found a positi\.e relationship
t ) c , ( \ \ (YW ;ts\-mnietry in the numhcr of iintlcr / J O ~ ~ and
I ~ S thc total numher of point5
1 1 1 1 \hi%l a q t - s t mtler. I\"c should note. IioLveiw, that this rcsult is espi~rsscdin ;t
v )ni[s\vliat dil'frrent currency than other reported rcsults on simple antler Icngtli in
t l i a t it chis Lvitli complexity rather than antler size. Furthermorc, these authors did
h l i o \ \ , that the precise form of the relationship bet\\-een as):mnictr>-in aiitlcr points
a l i t l total ~iunil~eiof points on thr largest aiitler revcaled a significant age-dcpendcni
clt%cwasc iii rcgrcssion c.oefficients, suggesting a greater teiideiic>. t o 11iitfi.r dc\ c~lopnictitalas~-mmcti-y
a n m i g older bulls (Sollm-g 8r Saetlirr, 1993).
lr \vould thus appear that \\.hatever ma)- he the environmental stresses o f
t l ~ ~ ~ . c ~ l i ~ p idi idcc~ri iniales
i,
of all species of deer so far studied niatiagc t o achici-t.
1111)rt. syntiit~trical antlcrs, in ternis of o\-erall antler length, t h a n do >.ouiigt*r
in(li\,~iciual~.
Such conclusion is at least not inconsistent with thc idra that fkrnales
iiiii). uw aiitlvr s!ninietr>- a s an iinambiguous indicator of fitnrss, as sugg:c*strtlniortq:c~ic.rall\1)). Illoller & H~g1;luiid( 199 1). hloller (1992, 1993).
.Is the numher of studies of FL4continucs to expand. it i h becoining appar
t1i;it F.1 ma\- offer a po\verful tool for distinguishing lict\veeii iilternati\.c h \ ~ o t h c
o f tlie intcxtctions of varied selcction pressures on development and morpholoq.
\\'(. predict that studies such as those of Rloller 6: Poniiankoc\&i ( 1993ti'i. u,hich u s c
F.1 I o di4nguish bct\veen sexual traits subject to Fisheriaii o r Zaha\.ian ~ l e c t i o i i .
; i n t i die aiialysis presentcd here, which attempts to use FA to identify the age a t
\I Iiic h sc.sual sclcction for symmetrical ornaments can out\\~eigh thc Yffkcts i )f
dc.\.c~lol~iii~.ii\al
stress, [\-ill pro\^ to lie an important use for fiiturc anal>.sis of FA\.
Consenation Trust who a s s i s t d with many of the catching exercises. XI1 l i v e
capture operations wrrc undertakcri undcr appropriate licence fi-om English Naturr.
Finally \ve would thank John Allen and t\vo anonymous rcfcrees for constructive
comments on an earlier draft of this paper.
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