American Fisheries Society Symposium 23:77–87, 1999

© Copyright by the American Fisheries Society 1999

Application of Radiometric Age Determination to Three

Long-Lived Fishes Using 210Pb:226Ra Disequilibria in
Calcified Structures: A Review

ERICA J. BURTON, ALLEN H. ANDREWS, KENNETH H. COALE,

AND GREGOR M. CAILLIET
Moss Landing Marine Laboratories
Post Office Box 450, Moss Landing, California 95039-0450, USA

Abstract.—Radiometric aging of fishes is a recently developed technique using the disequi-

libria of 210Pb:226Ra in calcified structures to determine age. It has been applied successfully
to several fish species, but certain limitations have made improvements desirable. Because
226
Ra can be measured directly by counting atoms using isotope-dilution thermal ioniza-
tion mass spectrometry (TIMS), a new ion-exchange separation technique was developed
to isolate small quantities of radium from calcified structures for TIMS determination. The
advantages of this new technique are reduced sample mass and processing time, and greater
accuracy and precision of radium quantification. We applied this technique to calcified
structures from three fish species: otolith cores of Pacific grenadier Coryphaenoides acro-
lepis and tarpon Megalops atlanticus, and pectoral fin ray cores of Atlantic sturgeon Acipen-
ser oxyrinchus. Annulus-derived age estimates for C. acrolepis were accurate with a
confirmed longevity of at least 48 years. Although annulus-derived ages for M. atlanticus
were inconsistent with radiometric ages, radiometric aging confirmed that tarpon are long-
lived; females may exceed 82 years. Radiometric age could not be determined for A.
oxyrinchus because 210Pb activities were greater than could be supported by ingrowth from
226
Ra decay. In this paper we discuss the application and limitations of this technique and
its relevance to fisheries management.

Traditional age validation methods (mark-and- techniques to long-lived or difficult-to-age fishes.

recapture, laboratory rearing, or marginal incre-
ment analysis) are often difficult or impractical
to use for long-lived or deep-sea fishes. Fortu-
nately, a naturally occurring radioisotope pair,
radium-226 (226Ra) and its daughter product lead-
210 (210Pb), found in calcified structures of fishes
can act as a built-in chronometer. This radioiso-
tope pair is useful for measuring age in time
scales up to about 120 years.
Radiometric age determination of fishes is
most useful for aging difficult-to-age or difficult-
to-sample species. The technique cannot rou-
tinely be used for all aging studies, because it is
costly and few laboratories have the specialized
equipment necessary to process and age calci-
fied structures.
At least fifteen papers have been written on
radiometric age determination of fishes, since
Bennett et al. (1982) confirmed the longevity of
the splitnose rockfish (Sebastes diploproa) using
210
Pb: 226Ra disequilibria in otoliths ( Table 1).
Eleven of these studies attempted to apply these
Two of the eleven studies were unable to confirm
age estimates, because three fundamental radio-
metric assumptions may not have been met
(Welden et al. 1987; Fenton et al. 1990). Nine of
the eleven studies successfully confirmed otolith
age estimates or longevity (Campana et al. 1990;
Fenton et al. 1991; Watters 1993; Kastelle et al.
1994; Fenton and Short 1995; Milton et al. 1995;
Stewart et al. 1995; Kline 1996; Andrews 1997).
Previous studies that used 210Pb:226Ra dis-
equilibria to determine fish age measured 226Ra
activity either indirectly, using the decay of ra-
don-222 as a proxy for 226Ra (radon emanation;
Bennett et al. 1982; Campana et al. 1990; Watters
1993; Kastelle et al. 1994; Kline 1996); or directly,
using the alpha-decay of 226Ra (␣-spectrometry;
Fenton et al. 1990; Fenton et al. 1991; Milton et
al. 1995; Stewart et al. 1995). These two methods
typically require 1–10 g of calcified material, 4–6
weeks processing time, and can result in high
analytical uncertainty (4–150%). Improvements
to the existing methods were desirable, includ-

77
78 BURTON ET AL.

TABLE 1. Radiometric age determination, review, and modeling papers published since 1982. Reference,
subject of paper, radionuclide pair used (or discussed), species examined, common name, structure used
(or discussed) for radiometric analysis, results, and useful aging technique (if applicable) are noted.

Reference Radionuclide Species Structure used Results

(Subject of paper) pair (common name) for radiometry (aging technique)

210
11. Bennett et al. 1982
(age determination)
210
12. Welden et al. 1987
(age determination)
Pb:226Ra Sebastes diploproa Otoliths Confirmed longevity
(splitnose rockfish) (whole) about 80 yr (otolith
sections)
Pb only Triakis semifasciata Cartilaginous Unsuccessful age
(leopard shark) vertebrae determination for all
four species;
suggested closed
system assumption
Alopias vulpinus violated
(thresher shark)

Squatina californica
(angel shark)

Carcharodon
carcharias
(white shark)
210
13. Campana et al. 1990 Pb:226Ra Sebastes mentella Otoliths Confirmed age
(age determination) (deepwater redfish) (cored) estimates up to 65 yr
(break-and-burn)
210
14. Fenton et al. 1990 Pb:226Ra Macruronus Otoliths Unsuccessful age
(age determination) novaezelandiae (whole) determination;
(blue grenadier) suggested constant
uptake assumption
violated
210
15. Fenton et al. 1991 Pb:226Ra Hoplostethus atlanticus Otoliths Confirmed long-lived
(age determination) (orange roughy) (whole) (about 149 yr)
228
16. Campana et al. 1993 Th:228Ra Hirundichthys affinis Otoliths Confirmed age
(age determination) (fourwing flyingfish) (whole & cored) estimates and
longevity less than 2
yr (otolith volume &
known-age fish)
210
17. Watters 1993 Pb:226Ra Sebastes rufus Otoliths Confirmed age
(age determination) (bank rockfish) (cored) estimates up to 40 yr
(break-and-burn &
sections)
210
18. Kastelle et al. 1994 Pb:226Ra Anoplopoma fimbria Otoliths Confirmed age
(age determination) (sablefish) (cored) estimates up to 23 yr
(whole & break-and-
burn)
210
19. Fenton and Short 1995 Pb:226Ra Macruronus Otoliths Approximated age
(age determination) novaezelandiae (cored) estimates up to 13 yr
(blue grenadier) (whole & sections)
210
10. Francis 1995 Pb:226Ra Hoplostethus atlanticus Otoliths Re-confirmed long-lived
(modeling & (orange roughy) (whole) over 100 yr
reanalysis of
published data)
 RADIOMETRIC AGE DETERMINATION OF LONG-LIVED FISHES 79

TABLE 1. (continued)

Reference Radionuclide Species Structure used Results

(Subject of paper) pair
210
11. Kimura and Kastelle
1995 (modeling,
review, and
recalculation of age)
228
210
12. Milton et al. 1995
(age determination)
(common name) for radiometry (aging technique)
Pb:226Ra Various Otoliths Discussed importance
(whole & cored) of using cores
Th:228Ra Hirundichthys affinis (whole & cored)
(fourwing flyingfish)
Pb:226Ra Lutjanus erythropterus Otoliths Confirmed whole
(Crimson snapper)a (whole & cored) otoliths better
predictor than
sections for all
three species

Lutjanus malabaricus Otoliths

(Malabar snapper)a (whole & cored)

Lutjanus sebae Otoliths

(Emperor snapper)a (whole & cored)
210
13. Smith et al. 1995 Pb:226Ra Hoplostethus atlanticus Otoliths Confirmed long-lived
(reassessment of (orange roughy) (cored) up to 101–129 yr;
radiometric age) Maximum age
converges with
previous estimates
210
14. Stewart et al. 1995 Pb:226Ra Allocyttus verrucosus Otoliths Approximated age
(age determination) (warty oreo) (whole) estimates; Confirmed
long-lived up to 132
yr (otolith sections)
210
15. Kline 1996 Pb:226Ra Sebastolobus alascanus Otoliths Confirmed age
(age determination) (shortspine (cored) estimates up to 80 yr
thornyhead) (otolith sections)

Sebastolobus altivelis Otoliths Confirmed age

(longspine (cored) estimates up to 45 yr
thornyhead) (otolith sections)
210
16. Andrews 1997 Pb:226Ra Coryphaenoides Otoliths Confirmed age
(age determination) acrolepis (whole & cored) estimates up to 56 yr
(Pacific grenadier) (otolith sections)

ing increased accuracy and precision of radium Radiometric Assumptions

measurements and analyzing single otoliths
rather than pooling otoliths into age groups.
Because 226Ra can be measured directly by
counting atoms on a thermal ionization mass
spectrometer, a new ion-exchange separation
technique was developed to isolate small
quantities of radium from calcified structures
(Andrews 1997). The advantages of this new
technique are reduced sample mass and pro-
cessing time, and greater accuracy and preci-
sion of radium quantification.
Radiometric age determination of fishes re-
lies on the incorporation of 226Ra (a calcium ana-
log) from the environment into the calcified
matrix of otoliths and skeletal structures. Once
226
Ra is incorporated into the structure, it decays
through a series of short-lived daughter isotopes
to the more stable isotope 210Pb. In an ideal situ-
ation, 210Pb found in calcified structures exists
only as ingrowth from incorporated 226Ra, where-
as 210Pb is not incorporated into closed calcified
systems from the surrounding environment; and
80 BURTON ET AL.
226
Ra and its daughters do not migrate. By mea-
suring both 210Pb and 226Ra, the time that has
passed since the initial uptake of 226Ra can be
determined (i.e., since birth).
There are three assumptions associated with
radiometric age determination. First, the calci-
fied structure must act as a closed system for ra-
dium and its daughter products. Violation of a
closed system is unlikely in otoliths, because it
consists of a well regulated aragonitic lattice
where crystalline growth is directionally regu-
lated by polyanions (Wheeler and Sikes 1984),
and otolith resorption appears to occur in very
rare cases or not at all (Campana 1983; Ichii and
Mugiya 1983; Yoklavich and Boehlert 1987). In
addition, a significant loss has not been docu-
mented in less controlled forms of calcium car-
bonate, such as corals (Bender 1973; Moore et al.
1973; Dodge and Thompson 1974). We did not
investigate this assumption, because numerous
other studies have demonstrated the successful
application of radiometric age determination.
Second, the uptake of 210Pb is either very low
or insignificant during the formation of the calci-
fied structure, or at least in the core (the first few
years of growth). This initial uptake of 210Pb is ide-
ally close to zero and inferred from young fish. We
investigated this assumption by measuring 210Pb
and 226Ra in calcified structures of young-of-the-
year or juvenile fish. Because age is less question-
able for young fish, age was determined using
210
Pb:226Ra ratio. Effectively, age estimates would
be significantly greater than expected if 210Pb is
incorporated at significant levels.
Third, the uptake of 226Ra is proportional to
mass growth of the calcified structure. This as-
sumption is only necessary when whole struc-
tures are used and is largely circumvented by
using cored structures (Campana et al. 1990;
Kimura and Kastelle 1995). In our studies, cores
of otoliths or pectoral fin rays were used and
210
Pb:226Ra was measured for each sample.
Radium Separation
As calcium is incorporated into the calci-
fied structures of fish, barium and radium are
also incorporated because they are calcium ana-
logs. Calcium and barium are present in much
greater quantities than radium (Dannevig 1956;
Macpherson and Manriquez 1977; Edmonds et
al. 1991). These elements can suppress the ra-
dium signal when analyzed using TIMS; there-
fore, they must be significantly reduced in the
sample prior to performing isotope-dilution
TIMS.
To isolate small quantities of radium from
calcium and barium, a new ion-exchange sepa-
ration technique was developed (Andrews 1997).
This technique significantly reduces calcium and
barium in the sample, while conserving radium,
using a series of cation-exchange columns. The
technique was modified from an existing geologi-
cal technique used for isolating radium from vol-
canic rocks (Chabaux et al. 1994).
There are several advantages to using the
new radium separation and TIMS detection
techniques. First, the thermal ionization mass
spectrometer counts radium atoms directly,
rather than relying on activity or decay prod-
ucts. Second, only 0.1–1.0 g of calcified mate-
rial is required, depending on 226Ra activity.
Third, the analytical uncertainty (counting er-
ror) using TIMS is usually less than 1.5%. Finally,
the radium processing time is reduced from 4
to 6 weeks to 7–10 d. In summary, this modifi-
cation to the radiometric aging technique
requires less calcified material, reduced pro-
cessing time, and radium quantification is more
accurate and precise.
In three concurrent studies, the new radium
separation technique was applied to three diffi-
cult-to-age fish species: Pacific grenadier Cory-
phaenoides acrolepis, tarpon Megalops atlanticus,
and Atlantic sturgeon Acipenser oxyrinchus. The
Pacific grenadier is a deep-sea species for which
there are conflicting age and longevity estimates,
ranging from 6 to greater than 60 years (Kulikova
1957; Mulcahey et al. 1979; Wilson 1982; Matsui
et al. 1990). The tarpon is a large, migratory fish
that occupies a variety of tropical and sub-tropi-
cal habitats (Zale and Merrifield 1989). It is sug-
gested from otolith annuli that the tarpon is
long-lived, at least 55 years (Crabtree et al. 1995).
The Atlantic sturgeon, an anadromous fish, occurs
in fresh, estuarine, and coastal waters. It is sug-
gested from annuli found in pectoral fin rays that
the Atlantic sturgeon is a long-lived species; age
estimates exceed 40 years. In this paper, we
present three radiometric aging studies with dif-
fering conclusions: (1) Radiometric confirmation
of Pacific grenadier annulus-derived age estimates
and longevity; (2) Radiometric confirmation of
tarpon longevity; and (3) Possible violation of the
closed system assumption in pectoral fin rays of
Atlantic sturgeon.
 RADIOMETRIC AGE DETERMINATION OF LONG-LIVED FISHES
Methods
Sample Preparation
Pacific grenadier.—To estimate age, otoliths
were transversely sectioned and annuli quanti-
fied. Based on annulus quantification, otolith
mass, capture date, capture location, and sex,
otoliths were pooled into seven age groups for
radiometric analysis. Pooled otoliths were cored
to the dimensions and mass of a 5-year-old
otolith. The pooled otolith cores were processed
using the modified radiometric aging technique
(Andrews 1997).
Tarpon.—We obtained whole otoliths of tar-
pon from the Florida Department of Environ-
mental Protection, which had been aged by
counting annuli in transverse sections (R. E.
Crabtree, Florida Marine Research Institute, per-
sonal communication). We attempted to confirm
annulus-derived age estimates using the modi-
fied radiometric aging technique.
Otoliths were cored to the size of a 2-year-
old otolith, and pooled into age groups based on
capture date, sex, and annulus-derived age
(Andrews et al. 1997b). Otoliths were analyzed
from six pooled age groups. In addition, we ana-
lyzed nine single otolith cores in an attempt to
age individual fish using the radiometric aging
technique. Because the size of a single 2-year-old
otolith core was relatively large (approx. 0.1 g),
and TIMS is very sensitive, single core analysis
was possible. Radiometric methods were then
applied to the tarpon (Andrews et al. 1997b).
Atlantic sturgeon.—We obtained pectoral fin
rays of Atlantic sturgeon from the Chesapeake
Biological Laboratory, which had been aged from
transverse sections, providing age estimates from
5 to 42 years (D. H. Secor, University of Maryland,
personal communication).
Use of sturgeon otoliths was not feasible
because they are very small, irregularly formed,
and there is a moratorium on fishing, making
new specimens difficult to obtain. Because fin
rays are large and form regular annuli, we were
able to analyze single fin rays and pooling of age
groups was unnecessary.
In this study, we examined two whole juve-
nile pectoral fin rays (estimated age of 5 years),
and four cored adult fin rays (first 5 years of fin
ray growth). Radiometric age determination was
applied to these samples (Andrews et al. 1997a).
81
Radiometric Methods
Trace-metal clean procedures and equip-
ment were used throughout the analyses to avoid
contamination of lead, barium, and calcium
(typically found on laboratory glassware). All ac-
ids were double distilled (GFS Chemicals®) and
dilutions were made using Millipore® filtered
Milli-Q water (18 M⍀ cm–1). Calcified structures
were cleaned according to procedures described
in detail elsewhere (Andrews 1997). Polonium
was separated from samples by autodeposition
onto a silver planchet (Flynn 1968). Activity of
210
Pb was determined using ␣-spectrometry of
210
Po, its daughter product. Radium-226 was
separated from calcium and barium using a se-
ries of cation-exchange columns, and radium
was quantified using TIMS (Andrews 1997).
Results
Pacific Grenadier
Annulus-derived ages and radiometric ages
were in close agreement (Figure 1). There was no
significant difference between the two age esti-
mates (paired two-tail t-test, df = 6, t = 0.4269,
P = 0.6844). According to annulus-derived age,
the youngest age-group was 1–3 years, which was
radiometrically aged as 0.6 year. The oldest an-
nulus-derived age-group, composed of males,
was estimated to be 46–56 years; radiometric age
was determined as 56 years.
Tarpon
Annulus-derived ages and radiometric ages
were not in close agreement (Figure 2). Analyti-
cal uncertainty of radiometric measurements for
a couple of samples, however, coincided with the
line of agreement. Although there was no signifi-
cant difference between the two age estimates
(paired two-tailed t-test, df = 14, t = 0.4181, P =
0.6822), residuals were large, the coefficient of
determination was low (adjusted r 2 = 0.552;
Kvålseth 1985), and the power to detect a 5-year
difference was low (1 – ␤ = 0.74).
Age estimation seemed to differ between age
groups and the sexes (Figure 2). Male annulus-
derived age estimates were greater and less than
radiometric ages with no apparent pattern asso-
ciated with age. In contrast, female annulus-
derived age estimates were consistently greater
than radiometric ages for fish less than 40 years
old (radiometric age), but not for one sample older
than that. The oldest female annulus-derived age
82 BURTON ET AL.

FIGURE 1. Comparison of male, female, and immature Pacific grenadier Coryphaenoides acrolepis mean
annulus-derived ages and radiometric ages. A linear regression and a line of agreement are drawn for
comparison (adjusted r2 of regression noted; Kvålseth 1985). Vertical bars represent low and high radio-
metric age estimates, based on analytical uncertainty of 210Pb and 226Ra measurements. Horizontal bars
represent the range of annulus-derived ages for each pooled otolith age group. (Source: Andrews 1997)

estimate (55 years) was underestimated by almost Discussion

30 years (radiometric age; 82 years). The upper
ratio estimate for this sample exceeded a ratio of
1.0; which makes the upper radiometric age esti-
mate undefined (Figure 2). This may be attributed
to the small sample size (approximately 0.09 g)
and low 210Po activity; which approaches the de-
tection limits of the ␣-spectrometer.
Atlantic Sturgeon
Radium activity was fairly consistent among
pectoral fin ray samples (0.068 ± 0.012 disintegra-
tions per minute per gram; Andrews et al. 1997a).
Unfortunately, 210Pb activities were greater than
could be supported by ingrowth from 226Ra decay;
210
Pb activities were 5–41 times greater than ex-
pected. The levels of exogenous 210Pb were vari-
able among samples, and a correction could not
be made. Therefore, age determination was not
possible.
Pacific Grenadier
Annuli found in otolith transverse sections
can be useful and valid for aging Pacific grena-
dier. Although the oldest age-group was validated
as 56 years, the greatest annulus-derived age es-
timate was for a 73-year-old male. Females are
also long-lived, but as the number of annuli in-
crease, otoliths become more difficult to age.
Radiometric age determination of fish older than
about 56 years could not be performed because
of low sample availability.
Historic and recent age estimates for the
Pacific grenadier are controversial and contra-
dictory. Kulikova (1957) estimated a longevity of
6 years using scales. Mulcahey et al. (1979)
found otolith transverse sections difficult to age
and cautiously estimated age as 15–25 years.
Wilson (1982) examined scales, vertebrae, and
 RADIOMETRIC AGE DETERMINATION OF LONG-LIVED FISHES 83

FIGURE 2. Comparison of male and female tarpon Megalops atlanticus annulus-derived ages (mean age,
if otoliths pooled) and radiometric ages. A linear regression and a line of agreement are drawn for com-
parison (adjusted r2 of regression noted; Kvålseth 1985). Vertical bars represent low and high radiometric
age estimates, based on analytical uncertainty of 210Pb and 226Ra measurements. Horizontal bars represent
the range of annulus-derived ages for each pooled otolith age group (if applicable). The upper limit of the
radiometric age estimate for the oldest female is undefined, because analytical uncertainty of 210Pb: 226Ra
exceeds 1.0. (Source: Andrews et al. 1997b)

otoliths, and estimated a longevity of approxi- Tarpon

mately 20 years. Matsui et al. (1990) also had
difficulty reading otoliths, especially near the
periphery of otoliths from large C. acrolepis in-
dividuals. Longevity estimates from Matsui et
al. (1990; 62 years for males and females) closely
agree with radiometric age estimates for a simi-
lar size-class (56-year-old males, approximately
23 cm preanal fin length; Andrews 1997). Based
on annulus-derived ages, longevity may ap-
proach 73 years for male Pacific grenadier
(Andrews 1997).
A rapidly growing fishery exists off Califor-
nia and Oregon for the Pacific grenadier (R. Leos,
California Department of Fish and Game, per-
sonal communication). Based on annulus-
derived ages and radiometric ages, the Pacific
grenadier are long-lived. Therefore, conservative
measures need to be taken in this fishery.
Our study of tarpon presents the first radio-
metric age determination ever performed on in-
dividual otolith cores, and consequently, on
individual fish. Single otolith analysis was pos-
sible because of the large size of a 2-year-old
otolith core (approximately 0.1 g), relatively high
226
Ra activity (0.0435–0.3951; N = 9), low detec-
tion limits and great analytical precision of the
new TIMS technique (<1.5% standard error at
95% confidence level).
Although a significant difference between
radiometric and annulus-derived age estimates
was not detected, the linear regression charac-
teristics suggested a statistical test was suspect;
residuals were large, the coefficient of determi-
nation was low, and the power to detect a 5-year
difference was low. The two age estimates may
be biologically different (Yoccoz 1991) and could
84 BURTON ET AL.
have an affect on population parameters, man-
agement, or conservation policies.
Annuli found in otolith transverse sections
may not be useful for aging tarpon. Tarpon
otoliths are difficult to read and can be confus-
ing (Crabtree et al. 1995). The difficulty in read-
ing otoliths, and the error associated with
assigning an age from multiple readings, may
have been why annulus-derived ages were not
in close agreement with radiometric ages.
Annual periodicity of growth increments has
been validated using oxytetracycline (OTC)
markers for twelve of eighteen captive tarpon,
ranging in age from 4 to 9 years (Crabtree et al.
1995). Marginal increment analysis of young-of-
the-year and 1-year-old tarpon otoliths also sup-
ported the hypothesis that growth increments
formed annually. It is uncertain, however, if this
trend continues in the adult life history stages.
Because this study used single otolith cores,
did not produce 210Pb:226Ra ratios greater than 1.0,
and did not produce ages that were greatly dif-
ferent from annulus-derived ages, radiometric
age estimates may be more accurate than annu-
lus-derived age estimates. Radiochemistry con-
firmed that tarpon are long-lived and attests to
the difficulty of aging tarpon by quantifying an-
nuli. It seems appropriate that radiometric age
determination is a necessary component in age
determination of the tarpon.
In southern Florida and parts of Central
America, tarpon are the basis of economically
important recreational fisheries (Crabtree et al.
1995). In Florida, the sport fishery is intensely
regulated. Since the establishment of a permit
system in 1989, the harvest of tarpon has de-
clined to less than one hundred fish per year, and
the fishery is now mostly catch-and-release. The
confirmed longevity of the tarpon is a strong ar-
gument for sustained and geographically ex-
panded regulations.
Atlantic Sturgeon
Pectoral fin rays showed signs of vascular-
ization in the core region that extended through
the fin rays. Brennan and Cailliet (1991) de-
scribed pectoral fin ray sections of the white stur-
geon as having a “marbled” appearance caused
by several morphological characteristics, includ-
ing numerous vascular channels. Core matrix
expansion and vascularization of dorsal and anal
fin spines have also been described for the At-
lantic sailfish ( Jolley 1977; Prince et al. 1986) and
several marlin species (Prince et al. 1984; Hill et
al. 1989). Hoenig and Walsh (1982) noted the
presence of vascularized canals in vertebrae of
several elasmobranch species, a morphological
characteristic that may have violated the closed
system assumption in an age determination
study of several shark species conducted by
Welden et al. (1987). Vascularization and core
matrix expansion throughout the fin ray of At-
lantic sturgeon may allow nuclide migration.
Therefore, the closed system assumption may
have been violated. The successful application
of the radiometric method to Atlantic sturgeon
will require that closed calcified systems, such
as otoliths, be used.
Although Atlantic sturgeon age estimates
could not be confirmed with radiochemistry
(Andrews et al. 1997a), annual growth zone for-
mation was confirmed using marginal increment
analysis, OTC-marking, and laboratory rearing
(Secor et al. 1997). Annuli found in pectoral fin
rays of other sturgeon species have also been
validated with tags and OTC markers (Acipenser
transmontanus, Brennan and Cailliet 1991; Aci-
penser fulvescens, Rossiter et al. 1995). All of these
studies suggest sturgeon are long-lived.
Until 1996, the Hudson River’s Atlantic stur-
geon population supported a commercial fish-
ery. A moratorium, however, currently exists on
Atlantic sturgeon harvest (Secor et al. 1997).
Based on longevity estimates, there is little doubt
that this slow-growing, long-lived species will
require many years to recover.
Conclusions
In summary, the new radium separation
technique coupled with isotope-dilution TIMS,
has allowed us to use smaller amounts of calci-
fied material in the radiometric aging process.
This has enabled us to age individual fish.
Pacific grenadier and tarpon are long-lived
species. Annuli found in Pacific grenadier otolith
transverse sections can be useful and valid to
determine age. Annuli found in tarpon otoliths,
however, appear to be unreliable for age deter-
mination. Therefore, the tarpon is an example of
a species where radiochemistry is an essential
component of age determination.
Although we can minimize or eliminate two
of the three radiometric aging assumptions by
using cores of calcified structures, the closed sys-
tem assumption may complicate analysis of sec-
ondarily calcified systems. Age validation for
chondrichthyans (sharks), chondrosteans (stur-
 RADIOMETRIC AGE DETERMINATION OF LONG-LIVED FISHES
geon & paddlefishes), and probably all fishes, will
need to use a combination of techniques, includ-
ing tag-recapture and marking techniques.
The radiometric aging technique is useful for
the group of fishes where the necessary assump-
tions are met for a given calcified structure, par-
ticularly the closed system assumption. This
technique has worked well for several species
using otoliths: 1) rockfishes (Bennett et al. 1982;
Campana et al. 1990; Watters 1993), 2) orange
roughy (Fenton et al. 1991; Smith et al. 1995), 3)
sablefish (Kastelle et al. 1994), 4) blue grenadier
(Fenton and Short 1995) and Pacific grenadier
(Andrews 1997), 5) several tropical snappers
(Milton et al. 1995), 6) warty oreo (Stewart et al.
1995), and 7) two thornyheads (Kline 1996).
Acknowledgments
We thank Bob Lauth (Northwest Fisheries
Science Center, National Marine Fisheries Ser-
vice [NMFS], National Oceanic and Atmospheric
Administration [NOAA]), Michael Hosie (Oregon
Department of Fish and Wildlife), and John But-
ler (Southwest Fisheries Science Center, NMFS,
NOAA) for shiptime and Pacific grenadier
samples; Roy Crabtree (Florida Department of En-
vironmental Protection, Florida Marine Research
Institute) for tarpon otoliths; David Secor and
Edward Houde (Chesapeake Biological Laborato-
ries at the University of Maryland) for pectoral fin
rays; Jocelyn Nowicki for assistance with develop-
ment of the radium separation technique and
alpha-spectrometry assistance; and Craig Lund-
strom, Xenon Palacz, and Peter Holden (Univer-
sity of California, Santa Cruz) for assistance with
development of the radium separation technique
and TIMS assistance. This paper was funded in
part by the Department of the Interior, U.S. Fish
and Wildlife Service, Federal Aid for Sportfish Res-
toration, Project Number F-59; the Hudson River
Foundation, Grant No. 006/93A; and the National
Sea Grant College Program, National Oceanic and
Atmospheric Administration, U.S. Department of
Commerce, under grant number NA36RG0537,
project number R/F-148, and under grant num-
ber NA89AA-D-SG138, project number R/ND-1-
20D through the California Sea Grant College
System, and in part by the California State Re-
sources Agency. The views expressed herein are
those of the authors and do not necessarily reflect
the views of NOAA or any of its subagencies. The
U.S. government is authorized to reproduce and
distribute for governmental purposes.
85
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