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The Forgotten Requirement for Age

Validation in Fisheries Biology
a

R. J. Beamish & G. A. McFarlane

Department of Fisheries and Oceans, Fisheries Research

Branch, Pacific Biological Station, Nanaimo, British Columbia,
V9R 5K6
Version of record first published: 09 Jan 2011.

To cite this article: R. J. Beamish & G. A. McFarlane (1983): The Forgotten Requirement for Age
Validation in Fisheries Biology, Transactions of the American Fisheries Society, 112:6, 735-743
To link to this article: http://dx.doi.org/10.1577/1548-8659(1983)112<735:TFRFAV>2.0.CO;2

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TRANSACTIONS

of the
November

AMERICAN

VOLUME

IISHERIE$ SOCIETY

NUMBER

1983
112

Tralsactlol,
oJthe
AmertcallFisherleSocle(3'
112:735 743, 1983
Copyrightby the AmericanFisheriesSociety1983

The Forgotten Requirement for Age

Validation in Fisheries Biology

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R. J. BEAMISHAND G. A. MCFARLANE
Department
ofFisheries
and Oceans,
Fisheries
Research
Branch
PacificBiological
Station,Nanaimo,BritishColumbiaV9R 5K6
Abstract

A surveyof 500 studiespublishedbetween1907 and 1980 that includedestimates

of fishage
indicatedthat only65% mentionedagevalidationor attemptedto validatethe ageingtechnique.
In lessthan 3% was the techniquevalidatedfor all age classes.Among the 35% that did not
consider
agevalidation,manydid not consider
the possibility
thatagesmaybe incorrect.Among
75 additionalpublications
publishedin primaryjournalsbetween1965 and 1980 that assessed
stockdynamicsand usedfishages,only40% mentionedor attemptedagevalidation,and none
successfully
validatedall agegroupsusedin the analysis.
Manyinvestigators
continueto neglect
the criticalstudyof age validationdespitethe cleardirectionof the early work on agedetermination.We showthat estimatedagesgreater than the maximumvalidatedage must not be
consideredaccurate.Use of inaccurateageshascausedseriouserrorsin the managementand
understanding
of fishpopulations.
Only by mark-recapturestudiesor useof known-age
fishcan
all ageclasses
in a populationbe validated.If suchstudiesare not possible,
fishshouldbe aged
by severalmethods,and the possibility
of errorsin ageestimatesmustbe considered.
ReceivedApril 5, 1983
Pioneer

work

on the determination

AcceptedAugust9, 1983
of fish

other populationsof the same speciesis dangerous.Validating a method of age determi-

agesstressedthe importanceof proving that a

technique was valid (Van Oosten 1923, 1929,

nation is as importantin fisheriesbiologyas

standardizingsolutionsor calibratinginstru-

1941; Hile 1936). In 1941, Van Oosten stated

that fisheriesinvestigatorshad "woefully ne- ments are in other sciences.

glected"to evaluatethe scalemethodof age
We believethe requirementto validateage
determination.He felt that thismethodof age determinationscontinuesto be neglected.To
determination
had been taken "too much for
examinethis, we reviewed500 papersthat use
grantedin thebeliefthat it isa simpleonewhich ageestimatesderivedfrom severaltechniques.
can be applied without technicalknowledge." The consequences
of not validatingagesare
He stressedthat the validityof a methodmust illustratedfor a commonfreshwaterspecies,
the
be establishedfor each speciesof fish studied: white sucker Catostomuscommersoni,and an im"It is a sadcommentaryon the attitudeof our portant marine species,the Pacificoceanperch
Sebastes alutus.

investigators
that althoughagedeterminations
of at least60 freshwaterspecieshavebeenpub-

lishedby more than 80 authorsin more than

150 papers,in only somehalf dozenpublicationsdid the authorsattempt to evaluatecriticallythe scalemethod."
All agesfor eachspeciesmust be validated.
Extrapolationbeyondthe maximum age validated, betweenspecies,or sometimesbetween

Terminologymust be standardized.Validationmeansprovinga techniqueis accurate.

Accuracycan be proven or estimated;estimatesof
accuracyare lessvaluable,but in some cases
only an estimateis possible.Precision
relatesto
reproducibilityand is not a measureof accu-

racy.The degreeof agreementamongreaders

is a measure of the precisionof the determi-

735

736

BEAM1SH AND MCFARLANE

TABLE1.--Numberofattempts
tovalidatemethods
ofage examinationof the edge of a structure, comdeterminations
from500 publications
thatincluded
age parison of back-calculated
lengthswith obestimates
for fish.
servedlengthsof the corresponding
agegroup,

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Publication

Degreeof

Pre- 1940- 1970-

validation

1940

1969

growth analysis,and comparisons

amongdifferenttechniques.
The maximumagevalidated

date

1980

Total

No validationattempted
Validationattempted

41
44

80
156

49
130

170
330

Validation

42

125

109

276

of first few

agegroupsonly
Validationof all agegroups

31

21

54

Validation

11

17

successful

was identified as well as the age range. It was

determinedif precisionwas estimatedand if

precisionwasusedto infer that the technique
wasaccurate.The studieswere separatedinto
three periods:pre-1940, 1940-1969, 19701980.

No age validationwasattemptedin 35% of

the 500 studies(Table 1). Most attemptsat valnationsand not the accuracyof the technique. idation involvedonly the first 1 or 2 years.In
An annulus,identified directly or indirectly,is 54 studies(9%), the techniqueusedwasapplianyzone,mark,or patternof growththatforms cableto all agegroups,but in only 17(3%)were
oncea year (Chugunova1959; Ricker 1968). all age groupsin the populationactuallyvaliThus, by definition,onlyoneannuluscanform dated. In almost all cases,validation of the first
or impliedto validate
in a year. These are the key termsrelevantto few yearswasconsidered
thisreport. Other termshavebeendefinedelse- for all age groups;that is, validation was extrapolatedbeyondthe oldestvalidatedage.Auwhere (for example,Chugunova1959).
thors of 29 paperscomparedmore than one
Literature Survey

structure. For 101 studies, it could be inter-

A surveyof 500 studiesthat involvedthe age- preted that precisionwas used to imply accuing of fish wasundertakento determinehow racy.
often, and to what extent, age-determination Validation was consideredin an increasing
methods were validated. Studies included in the
percentageof studiesin recentyears;however,
reviewwere publishedbetween1907 and 1980 the fraction of studiesthat involvedtechniques
in scientificjournals from severalcountries,the :apable of validating all age groups did not
majority(60%)havingbeenpublishedin North changein the two periodsfrom 1940 to 1980.
America. The papers were reviewed without Someauthors(27) referred to previousstudies
selectionand the numberof papersrelatedonly for validationof their techniques;of these,sevto the effort available.We doubt if increasing en referred to studies of Hile or Van Oosten
the samplesizewouldalter any conclusions.
In althoughboth theseauthorsstatedthat an agetechniqueshouldbe validatedfor
addition,we nonselectively
reviewed75 studies determination
in whichageestimates
wereusedspecifically
for populationsas well as for species.
Of the 75 papersthat dealt with stockdyanalysesof stock dynamics.These were publishedin fivemajorjournalsduring 1965-1980. namics, 40% mentioned or attempted validaA l'istof all reviewedpapersis availablefrom tion. Of these,none validatedagesfor all age
the atthors.
groupsusedin the analysis.
It issurprising,
perFor eachpaperit wasdeterminedif validation hapseven shocking,that 60% of thesestudies
of errors in
was attempted or mentioned. Validation was did not considerthe consequences
consideredsuccessfulwhen all reported ages age determination.A small percentage(8%)
were validated;that is, there were no agesre- evaluatedthe precisionof the age estimates.
The applicabilityof validationtechniquesin
portedthat wereolderthanthe oldestvalidated
age. For studiesin which validation was at- some of the studies in our review, or the relative
of somevalidationattempts,maybe subtempted,separationwasmadebetweenvalida- success
tion techniquesthat were capableof validating ject to differentinterpretations.However,such
agesfor all agegroupsand methodsthat apply differencesin interpretation will not alter the
only to the period of initial fastergrowth. The overwhelmingconclusion
that fisheriesbioloformer group includesmark-recapture tech- gistsseldomhavesuccessfully
validatedmethniquesand captureof known-agefish.The lat- odsusedto age fishdespitethe clear warnings
ter group includesan analysisof length-fre- of earlyworkers(VanOosten1923, 1929,1941;
quencymodes,monitoringof strongyearclasses, Hile 1936).

Downloaded by [171.67.34.205] at 07:15 18 March 2013

REQUIREMFNT
FORVAIIDATIONOFFISHA(;ES

737

1.0 mm

8615

8521

Fl(;Ure l .IScalesfromwhitesuckers
tagged
at age5 and recaptured
8yearslater.J = boundary
ofanteriorquadrazt;
B = posterior
edgeoJposterior
quadrant;J=focus.At thetbneoJrecapture
bothfish seemed
to beage5 bySpoor's
(I 939) criteria(axis_[-A).Olderagescoutdbeestima
ratedfrom irregula
riflesin theposterior
quadrant(axisf-B);
although
these
irregularities
produced
estimates
closer
totheknownageforfish8615, theydidnotdosoforfish8521.

Importance of Validation

Somebiologistsmay feel that we are overly

rigid in our insistence
that successlhl
validation
of an age-determination
methodmeansthat all
agegroupsbeingconsidered
are validated.Our
insistenceis basedon the consequences
of non-

White Suckers

The scale from the white sucker appears to

be a model tbr the scalemethod of age deter-

mination(Fig. 1 and Spoor 1939). Circuli are

distinctand widelyspacedduring periodsof ac-

tive fish growth;spacingbecomesreducedas

yearlygrowthdeclines.When scalegrowthreillustratethe importanceof suchconsequences. sumesin each year, the new circulus forms
around the incompletecirculi of the previous
In onecase,biologicalunderstanding
of a comyear. The annulusis definedas this zone of
mon freshwater species,white sucker, was
closelyspacedcirculi"cut off"'by the newcirimpeded for many years becauseonly young culus,and typicallyit is measuredwhere the
ageswere validated.In the other case,under- circulienter the posteriorquadrantof the scale.
estimatedagesof Pacificoceanperchmayhave In the anterior quadrantof the scale,the animposedmillion-dollarlosses
on the commercial nulusfrequentlyisassociated
with irregularities
in circuli formation.
fisheryfor that species.
validation,

and we offer two case histories to

738

BEAMISHAND MCFARLANE

Stewart(1926)validatedthe scalemethodby
PacificOceanPerch
followingtheyearlygrowthof scales
but only
The financialcostsand managementconsefor veryyoungwhitesuckers.
His application quencesof underestimatingthe ageof older fish
of the scalemethodshowed
thatfishgrewrap- can be illustrated for the Pacific ocean perch
idly and that growthincreasedin the 6th and

fisheryoff the westcoastof Canada.Prior to

7th years(webelievethisincrease
in growth 1980,fishin thecommercialcatchwerethought
resultsfrom underestimating
the ageof older
to be considerably
youngerthan nowbelieved.

fish).Spoor(1939)alsovalidatedthescalemethBiologistsof one of the major foreign nations

od for youngfish by examiningthe annual
fishingthis speciesusedscalesto estimateage
growthon the edgeof the scale.The oldestfish
(Chikuniand Wakabayashi
1970)and they bein his studywas10 yearsand 94% were 3-6
lievedthesefishwere rapid-growingand relayearsold. Eddyand Carlander(1942) examined
2,655 specimens
from 69 lakes;the oldestfish tively short-lived.Use of otolith surfacesby

North American scientists(Westrheim 1973)

indicatedthe fishwereolderthansuggested
by
temptedno validationbecause"for practical
scalesbut not asold asnow believedby people
purposes
the scalemethodof growthdetermiwhohavestudiedotolithsections
for thisspecies
nationis recognized
as valid." Subsequently,and other rockfish (Beamish 1979; Archibald
there havebeen publishedand unpublishedet al. 1981; Bennett et al. 1982; Chilton and

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reported was 9 years old but the authors at-

studies in which the scale method was used with-

Beamish 1982; Archibald et al., 1983).

out validation,the applicabilityof the scale

New estimatesof age have greatlyreduced
methodwasquestioned,or scaleswere shown
the estimatedrate of naturalmortalityandproto be unreliableand lessusefulfor agedeterduceda more conservative
managementstratminationthan other structures(Raneyand
egy (Archibald et al. 1981; Archibald et al.,
Webster1942;Rawson
1942;Ovchynnyk
1965;
1983). If this strategyhad been appliedat the
Geen et al. 1966; Coble 1967; Beamishand Harbeginning of the fishery and some educated
vey1969;Beamish1973;QuinnandRoss1982).
guesses
madeaboutlong-termyields,it can be
Earlystudies
indicated
thatwhitesuckers
grew estimated that lost wholesale value to Canadian
relativelyquicklyto ages4-7 years,and that
industry,in 1981 Canadiandollars,has been
fewsurvivedbeyondage9 or 10. Annualmorabout4 milliondollarsannually.The total loss
talitycalculated
fromtheseageswashighafter
maturityeventhoughactivegrowthcontinued. in yieldfor the northeastPacificwouldbe considerablyhigher. While it is a little unfair to
Recent studies that validated and used the finmakethiscomparisonbecauseit isquestionable
ray method(Beamishand Harvey 1969; Beamwhetheror notthe earlyfisherycouldhavebeen
ish,unpublished
data)showed
thata relatively
largenumberof fishin unexploited
populationsbetter managedevenwith our presentknowlthe costsand consurviveafter increasesin length ceaseor be- edge,this examplesuggests
sequences
of not knowingthat older, slowercomeminimal.There is a periodof very reducedgrowth (Beamish1973) and fish can be growingfish are an important componentof

mucholderthanpreviously
thought.The oldest
age(not validated)for thisspecies
is 23 years
(Fig.2) whichmorethandoubles
the earlyestimatesof the life spanof thisspecies.
Our understanding
of whitesuckerbiology
changedwhenfin raysratherthanscales
were
usedfor agedetermination.All annuliare distinct on sectionsof fin rays(Fig. 2) whereas
annulion scalesare difficultto identifyafter
aboutage 5 (Fig. 1). With hindsightafforded
by validatedages,irregularitiesin circulistructure in the posteriorquadrantof scales
maybe
interpretedas annuli but they are difficultto
defineandtheymaynot formeveryyear(Fig.
1).

the commercial

catch. The

use of incorrect

growthratescanalsoleadto rather largeerrors

in estimatesof stockproduction(Ricker 1969).
As demonstratedin thesestudies,it may be
difficultto determineagefrom somestructures
after the growth rate changesand growth incrementson a structure are reduced. This pe-

riod of reducedgrowthoften occursafter maturity, canbe quitelong(Power 1978;Beamish

1979; Beamish and Chilton 1982; Chilton and

Beamish1982), and possiblyis important in

maintainingpopulations(Leamanand Beamish,
in press).It is illogicalto dismiss
ageingerrors
asnot beingimportantwhenthedegreeof error
hasnot beenassessed
by validation.

REQUIREMENT FOR VALIDATION OF FISH AGES

25

739

2o
15

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IO

0.25

0.25

mrn

mm

5
6

15

l0
9

I0
II
12

4-

15
5

0.2

mm

0.1$ mm

FGURF
2.Pectoral fin-ray sections
from whitesuckers.
(A) Fishaged23+ years.(B-D) Fish8615, agedat 5+ years
at thetimeoftaggzng
(May 1973)(B)and 13+ yearsat thetimeofrecapture
Orune1981)(C).Insert(D) .hows
reduction
in spacing
between
theannuliwithage.(Sections
arefrom different
fin rays,hence
theirmorphologies
differ.Thewhole

fin frombothsides
shouldhavebeensampled.)

74o

BFAMISH AND MCFARLANF

FIN-

'

RAY

OTC

SECTION

MARK

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O.3

mm

B
OTOLITH

CROSS

OTC

SECTION

MARK

0.2

mm

SECONDDORSAL SPINE
OTC

MARK

0.7

mm

FIGUR
3.--Structureusedfor agedetermination
that weremarkedbyoxytetrao'cline
(OTC) injections.
(A) Fin-ray
section
from a lingcodOphiodonelongatusat liber(
2 ,earsafterijection.(B) Brokensection
ofan otolith
from a

sablefish
Anoplopoma
fimbria
atliberty
3years
afterinjection.
(C)Second
dorsal
spine
froma spiny
dogfish
Squalus
acanthiasat libertyI yearafterinjection.

REQUIREMENT FOR VALIDATION OF FISH AGES

Recommended

Validation

Methods

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The validationof agesof olderfishrequires

eithera mark-recapturestudyor the identificationof known-agefishin the population.It
may be possibleto find some"natural" mark
on a structurethat identifiesa particularyear
class,but suchmarksappearto be very rare.

741

grow more slowlythan females.The importanceof strongyearclasses

alsocanbe masked.
There

are other

more

subtle features

of a

populationthat maynot be detectedif it is not

known that older fish exist. Different growth
ratesamongstocksmay not be detected.The
importanceof longevityor the mechanisms
by
For mostspecies,
at thistime, there islittle choice whicha particularpopulationsurvivesin its enbut to undertakea mark-recapturestudyfor vironmentmaynot be considered.Somepeople
validation.The disadvantages
relateto thetech- mayfeel that these"ecological"considerations
nicalproblemsof applyinga tag and the time areunimportantor lessimportantfor managing
requiredfor growthof the particularstructure. a population. However, becausepopulations
for
However,the studywill allowall agegroupsto havehadto survivein changingecosystems
be examined.
thousands
of years,life-historyparametersare
At the timeof tagging,scales
or fin rayscan adaptive. Adaptation is brought about by
be removedfor comparisonwith similarstruc- changesin life-historytraits suchasage at matures upon recapture of the fish. The effect of turity, rates of growth, the number of annual
taggingand removalof structureson growth reproductions,and fecundity.These traits are
should be monitored but even if there is some importantandmustbeclearlyunderstood
when
initialeffectit probablywill not interferewith the responseof a population to the extreme
a validationstudy.It isalsopossible
to usechem- stressplacedon it by a commercialfisheryis
icals to mark structures. Fluorochrome
labels
considered.
Accurateagesare the key to definare most commonlyused, especiallywhen a ing theseparameters.
structure cannot be removed at time of markWe encouragebiologists
to re-examinesome
beliefssuchasthe generalappliing.They are readilyavailable,
easilyadminis- longstanding
tered, and provide a distinct fluorescentmark cabilityof the scalemethod for age determion a varietyof structureswhen exposedto ul- nation (Lux 1971; Everhart and Young 1981),
travioletlight (for example,Fig. 3). Appropri- or the view that there is close or isometric reatedosages
for oxytetracycline
are discussed
by lationshipbetweensize(length)of fishand size
Beamishet al. (in press).
of hard parts throughout the life spanof the
fish (Lagler 1956). We believe that there has
Conclusions
beena tendencyto forget the needto validate
In general,the age-determinationmethodse- age estimates,and that this may have led to
lectedfor manystudiesprobablyprovidesac- ratherseriousmisunderstandings
of the biology
curateestimatesof ageover the rangeof more and populationdynamicsof some important
rapid growth. However, when growth is re- commercialspecies.
duced becauseof the sex of an individual, matEstimatesof ageare difficultto produceand
uration,foodlimitations,changesin behavior, are not unequivocal.Becausethey are fundaenvironmental conditions, or other cause,it is mental to an understandingof the biologyof
possible,even probable, that there will be a fishesandthe dynamics
of fishpopulations
it is
changein the appearanceof an annulusandthat essentialthat they be provenaccurate.
the methodof age determinationwill have to
Acknowledgments
bemodified.Onlybyvalidatingthemethodcan
it be proven that fish are not older than estiClaudiaHand andKathyBestcarriedout the
matedand that older fishare not an important literaturereview.Doris Chilton, ShayneMaccomponentof the population.
Lellan, and Dawn Barnesassisted
in the prepIf estimates
of ageof older fishare not vali- arationof figuresand the estimationof some
dated,it is possiblethat errorsare occurring. ages.BruceLeamanprovidedestimates
of yields
Such errors will not be random and will be biased of Pacificoceanperch and reviewedthe manutowardsyounger ages, resulting in an accu- script.EdwardTripple suppliedthe datahe colmulationof estimates
in the vicinityof the age lectedfrom the three taggedwhitesuckersthat
at whicha particulartechniqueor interpreta- wererecaptured8 yearsafter beingtagged.The
tion breaksdown.Mortalityestimates
then can fin usedin Fig. 2A wassuppliedby Barry Pabe overestimated,
particularlyfor malesif they quin, Saskatchewan
ResearchCouncil.

742

BEAMISH AND MCFARLANE

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