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An holistic approach to fish stock identification

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DOI: 10.1016/S0165-7836(99)00065-X

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 Fisheries Research 43 (1999) 35±44

An holistic approach to ®sh stock identi®cation

Gavin A. Begga,*, John R. Waldmanb
a
National Oceanic and Atmospheric Administration, National Marine Fisheries Service,
Northeast Fisheries Science Center, 166 Water Street, Woods Hole MA 02543, USA
b
Hudson River Foundation for Science and Environmental Research, 40 West 20th Street,
9th Floor, New York, NY 10011, USA

Abstract

The concept of the `stock' is fundamental for both ®sheries and endangered species management. We review different
approaches used in identifying and classifying stocks and advocate that an holistic approach (e.g., involving a broad spectrum
of complementary techniques) be used in future stock identi®cation studies. Stock de®nitions should evolve as management
requirements do and be re-evaluated when the need arises or when break-through technologies become available. # 1999
Elsevier Science B.V. All rights reserved.

Keywords: Stock; Identi®cation; Population; Discrimination; Integration

1. Introduction problems in ®sheries management, and discuss the

The concept of the `stock' is fundamental to ®sh-
eries management. Stocks are arbitrary groups of ®sh
large enough to be essentially self-reproducing, with
members of each group having similar life history
characteristics (Hilborn and Walters, 1992). To man-
age a ®shery effectively, it is important to understand
the stock structure of a species and how ®shing effort
and mortality are distributed (Grimes et al., 1987). An
understanding of stock structure is vital to designing
appropriate management regulations in ®sheries
where multiple stocks are differentially exploited
(Ricker, 1981). We offer a review and perspective
on different approaches to ®sh stock identi®cation
*
Corresponding author. Present address: Marine Research
Institute, P.O. Box 1390, 121 Reykjavik, Iceland.
E-mail address: gavin@hafro.is (G.A. Begg)
utility of an holistic (multiple technique) approach as a
solution to these problems.
In ®sheries science, `stock' ®rst referred to any
group of a ®sh species that was available for exploita-
tion in a given area (Milton and Shaklee, 1987). Marr
(1957) explicitly differentiated stocks and subpopula-
tions, considering the subpopulation to be a geneti-
cally self-sustaining entity, i.e., a deme, or the smallest
self-perpetuating unit. He de®ned the stock as a
population or portion of a population, all members
of which are characterized by similarities which are
not heritable, but are induced by the environment, and
which may include members of several different sub-
populations. Race also has been used to describe
categories now considered to be stocks or populations
(e.g., Raney et al., 1954). Larkin (1972) emphasized
practical aspects and considered a stock to be a
production or management unit. Saila and Jones

0165-7836/99/$ ± see front matter # 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 1 6 5 - 7 8 3 6 ( 9 9 ) 0 0 0 6 5 - X
36 G.A. Begg, J.R. Waldman / Fisheries Research 43 (1999) 35±44
(1983) de®ned unit stocks as characteristic popula-
tions or sets of subpopulations within subareas of the
geographic range of a species, but noted that the
taxonomic status of unit stocks may vary or remain
unclear. However, a stock is often taken to mean a
lower category than that of the population recognized
by taxonomists (Cushing, 1968).
Modern stock de®nition studies often integrate
genetic knowledge, recognizing the importance of
information on the number and geographic limits of
non-interbreeding, self-recruiting populations within
an exploited species (Ovenden, 1990). In essence, the
stock concept describes the characteristics of the units
assumed homogeneous for particular management
purposes. There is a surprising degree of parallelism
between the problems in de®ning species (e.g., May-
den and Wood, 1995) and stocks: both have shifted
and diversi®ed over time partly in response to changes
in evolutionary thinking and technological advances.
Yet, on the operational level, the taxon levels of
species and stocks are usually differentiated without
debate. In contrast, the terms `stock' and `population'
are presently used rather interchangeably, although
there are exceptions. Regardless of its precise de®ni-
tion and biological underpinnings, the stock concept
really has to do with the interaction between a ®sh
species and its management. Consequently, the pur-
pose of such de®nitions is to direct management
efforts to taxon levels below that of the species. This
leads us to an important distinction to be made
between endangered species management and ®sh-
eries management with respect to stock identi®cation.
Endangered species management aims to ensure
that populations that exhibit unique life history traits
to particular areas are not irreversibly reduced by
harvest or habitat destruction (Dizon et al., 1992),
and is concerned with the evolutionary basis of stock
de®nition. Fisheries management, generally aims to
achieve maximum (or optimum) sustainable produc-
tion from ®sh stocks usually de®ned on the basis of the
same vital rate parameters that are used for produc-
tivity calculations. Therefore, the techniques used in
de®ning these two types of management needs are
fundamentally different. However, stock de®nitions
have tended not to be revisited in ®sheries manage-
ment because considerable ®scal and personnel
resources are normally required to conduct stock
identi®cation studies. Stock structure information
can often create more uncertainty in ®sheries manage-
ment, particularly when it contradicts historically
established stock and management boundaries, how-
ever, ignoring such information can contribute to
erroneous, ineffective ®sheries management.
There are many cases in which high exploitation,
combined with ineffective ®sheries management, have
resulted in depletion of ®sh stocks. These include
anchovy Engraulis ringens (Hilborn and Walters,
1992), capelin Mallotus villosus (Tjelmeland and
Bogstad, 1993), Atlantic cod Gadus morhua (Hutch-
ings, 1996), haddock Melanogrammus aegle®nus
(Clark et al., 1982), herring Clupea harengus (Dra-
gesund et al., 1980), orange roughy Hoplostethus
atlanticus (Smith et al., 1991), Atlantic salmon Salmo
salar (Cook and McGaw, 1991), and sardine Sardi-
nops sagax (Shannon et al., 1993). Such depletions
can result in a loss to the total gene pool of a species
(Nelson and SouleÂ, 1987; Smith et al., 1991), now a
consideration in the precautionary approach to ®sh-
eries management (ICES, 1997). Disregard of stock
structure in ®sheries management can also lead to
signi®cant changes in the biological characteristics of
a ®sh species (Altukhov, 1981; Ricker, 1981).
Identi®cation of ®sh stocks may be simple and
qualitative, such as observation of different timings
of runs of an anadromous species in a single river, or it
may be highly technical and quantitative, requiring
laboratory techniques and complex statistical ana-
lyses. Across this spectrum, scientists favor the former
approach for situations involving low numbers of
putative stocks exhibiting large characteristic inter-
stock differences, and favor the latter approach for
situations involving larger numbers of stocks and
where differences are slight between stocks.
The interplay of the particular questions posed,
resources available, and improvements in scienti®c
understanding and technology has resulted in a suite of
approaches applied to ®sh stock identi®cation. For
instance, in a review of the stock identi®cation history
of striped bass Morone saxatilis, Waldman et al.
(1988) found that stock identi®cation was based upon:
catch data, tag recoveries, meristics, morphometrics,
scale morphology, parasites, cytogenetics, protein
electrophoresis (isoelectric focusing), immunoge-
netics, mitochondrial DNA, and nuclear DNA. Heart
tissue fatty acids (Grahl-Nielsen and Mjaavatten,
1992), the elemental composition of otoliths (Mulli-
 G.A. Begg, J.R. Waldman / Fisheries Research 43 (1999) 35±44
gan et al., 1987), and osteological interdigitation
features (Waldman and Andreyko, 1993) have also
been used. Additional approaches such as stable iso-
tope measurements, and otolith microstructure, shape
analysis and thermal marking, have been applied to
other ®shes (e.g., Campana and Casselman, 1993;
Roelke and Cifuentes, 1997; Volk et al., 1999 these
proceedings).
Stocks are useful taxonomic groupings to the extent
that they are important for some explicit management
purpose. Many stock de®nitions used in ®sheries
management date back to when statistical data collec-
tion programs were ®rst implemented, and were often
based on geographical or statistical area boundaries
(e.g., in the northwest Atlantic, see Halliday and
Pinhorn, 1990). However, stock de®nitions should
evolve as management requirements do, and should
be revisited when the need arises or when break-
through technologies become available.
2. Fundamental `signals' from alternative stock
identification approaches
Fish stocks are identi®ed on the basis of differences
in characteristics between stocks. Quantitative bases
for demonstration of stock structure may occur as
discrete differences in some physical attribute, or as
a statistically signi®cant distribution. Investigation of
any single characteristic will not necessarily reveal
stock differences even when `true' stock differences
exist (Type 1 error). Characteristics vary between ®sh
stocks due to environmental and genetic factors.
Environmental factors tend to in¯uence phenotypic
characteristics within stocks; phenotypic variation
between stocks is usually associated with the geogra-
phical region occupied by a species throughout its
range. Although phenotypic differences do not pro-
vide direct evidence of genetic isolation between
stocks, they can indicate the prolonged separation
of postlarval ®sh in different environmental regimes
(Campana et al., 1995). A brief review underlying the
techniques for stock identi®cation follows:
2.1. Mark-recapture
Marking of ®sh can be used to infer stock structure.
For stock identi®cation purposes, best results occur
37
when putative stocks are marked when they are geo-
graphically discrete in order to determine whether
they subsequently intermingle. Alternatively, stock
mixtures may be marked to ®nd out if ®sh later
separate geographically. The success of mark-recap-
ture for stock identi®cation purposes is dependent on
representative tagging and recapture efforts (a condi-
tion rarely met). Such studies are generally costly and
time-consuming.
2.2. Catch data
Catch data can often be used as crude indicators of
stock structure. Strong geographic differences in age-
or size-composition, if not re¯ective of gear differ-
ences and other factors, suggest independence of
recruitment or other biological or ®shery factors as
a basis for assuming stocks are discrete.
2.3. Life history characteristics
Consistent differences in life history characteristics
such as growth rates or reproductive characteristics
(e.g., fecundity-at-age, spawning time, etc.) have fre-
quently been used to separate stocks.
2.4. Parasites
Parasite `tags' can also differentiate stocks of ®sh.
The species composition and abundance of parasites
may differ between ®sh stocks due to biogeography,
differential environmental tolerances of parasites, dif-
ferences in availability of intermediate hosts, and
different life history characteristics of the ®sh stocks
themselves. An advantage of this approach is that the
parasites are applied by nature at no cost to researchers
(Williams et al., 1992). Disadvantages include the
need for considerable biological information on the
parasites, and intra-stock variation due to adoption of
different life history modes within a ®sh stock (not to
mention the assumption that parasitic fauna are stable
over time, and the dif®culties in doing quantitative
parasitology).
2.5. Otolith microchemistry
Otoliths are predominantly composed of calcium
and trace elements that are derived from the ambient
38 G.A. Begg, J.R. Waldman / Fisheries Research 43 (1999) 35±44
waters inhabited by the ®sh (Campana et al., 1995).
Because water bodies often differ in the concentra-
tions of trace elements, stocks may often be distin-
guished by the chemical `signatures' retained in
otoliths (assayed using spectrometric or related tech-
niques). A further advantage of this approach is that by
analyzing selected portions of an otolith, the trace
element signals can be associated with particular
growth stages. This approach has potential to discri-
minate between stocks where an environmental signal
is pronounced (e.g., where substantial reproductive
interchange diminishes genetic differences). However,
results from these techniques are often dif®cult to
interpret because of the combined effects of physio-
logical, ontogenetic, and environmental in¯uences on
the deposition of trace elements, not to mention a
posteriori sampling problems with handling and ele-
mental contamination (Fowler et al., 1995; Thresher,
1999 these proceedings).
2.6. Morphology
2.6.1. Meristics
Countable, morphological structures (e.g., ®n rays,
gill rakers, scales in rows) have historically served as
an important basis for identifying ®sh stocks. Count
data are discrete, thus facilitating statistical analysis.
Meristics are controlled by both genetic and environ-
mental factors, in unknown proportions (Barlow,
1961). Meristic characters are generally set early in
ontogeny and remain stable throughout life; thus
re¯ecting environmental effects over a relatively brief
time of larval development. Because of this, signi®-
cant statistical differences can occur within a stock
among year classes or geographic subgroups subjected
to varying environmental conditions. However, con-
sistent environmental in¯uences have the potential to
provide stock discrimination where there is weak
genetic divergence between actual stocks.
2.6.2. Morphometrics
Morphometrics include the analysis of body shape,
or the shape of particular morphological features of
various body dimensions or parts. These data are
continuous, and must be corrected for size differences
among specimens. As with meristics, morphometric
expression is under the simultaneous control of
genetic and environmental factors.
2.6.3. Scale and otolith analyses
Geometric analysis of scales and otoliths provide
sources of variation amenable to morphometric and
other forms of analysis (e.g., Campana and Cassel-
man, 1993; Richards and Esteves, 1997). Features
containing stock-speci®c information such as annuli
spacing are biologically interpretable (i.e., related to
age and growth), whereas other features such as
perimeter shape are not easily interpretable.
2.7. Genetics
Genetic differences between individuals, stocks and
populations are the basis for ascertaining the degree of
reproductive isolation, which is the fundamental
mechanism structuring differences between these
taxonomic groups. The strengths of genetic signals
between stocks are positively associated with time
since divergence of stocks (mediated by generation
time, with shorter generation time accelerating genetic
differentiation), and their degree of isolation (i.e.,
reproductive exchange between stocks eroding
genetic differences) (Adkinson, 1996). Thus, recent
divergence, or substantial secondary reproductive con-
tact result in no apparent differences in gene frequen-
cies between groups, even when it exists. Genetic
techniques used in stock identi®cation include the
following:
2.7.1. Protein variation
Variation in proteins or enzymes is an indirect
expression of nucleotide base differences between
groups. Until the advent of molecular techniques,
allelic frequency differences revealed by protein elec-
trophoresis were the primary markers used to assess
genetic differences between stocks (Wirgin and Wald-
man, 1994). However, the usefulness of this approach
is undermined by the fact that the same protein may be
coded by multiple nucleotide sequences.
2.7.2. Mitochondrial DNA
Mitochondrial DNA (mtDNA) is a cytoplasmically
inherited circular molecule of approximately 16 000±
20 000 base pairs. Inheritance of mtDNA is haploid
and is either strictly or overwhelmingly maternal.
Different regions of the mtDNA molecule evolve at
different rates (Meyer, 1993), with differences in base
pairs used to separate stocks.
 G.A. Begg, J.R. Waldman / Fisheries Research 43 (1999) 35±44
2.7.3. Nuclear DNA
Base sequences of nuclear DNA (nDNA) parti-
tioned among chromosomes present large numbers
of potential markers (typically > 3 billion nucleotide
pairs) in any individual (Wirgin and Waldman, 1994).
Nuclear DNA is diploid; recombination re¯ects
genetic characteristics of both sexes. Nuclear DNA
that codes for gene products and functions is thought
to evolve substantially slower than mtDNA, but cer-
tain non-coding markers such as minisatellites and
microsatellites may evolve faster than mtDNA (Dowl-
ing et al., 1990). Interpretation of nDNA data may be
complicated by polyploidy, as is common in sturgeons
and salmonids (Wirgin and Waldman, 1994).
3. Integrated `signals' from multiple stock
identification approaches
The strongest inferences on stock structure are
drawn from a suite of complementary techniques that
cover multiple aspects of the biology of a ®sh species.
This is partly because the de®nition of a `stock', for
management purposes, is not strictly a genetic con-
struct, but represents a semi-discrete group of ®sh with
some de®nable attributes of interest to managers.
Integration of the results of each method used in a
multiple or `holistic' stock identi®cation approach
maximizes the likelihood of correctly de®ning stocks
± , however, de®ned by management (Hohn, 1997).
An holistic approach to ®sh stock identi®cation,
involving a broad spectrum of techniques, appears to
be particularly pertinent for those ®sh species with
complex stock identities. From an operational per-
spective we de®ne an `holistic approach' as one that
utilizes multiple techniques for ®sh stock identi®ca-
tion, and can include: collating all available stock
identi®cation information previously known into a
single review to infer stock structure; using two or
more different stock identi®cation techniques in a
single study on a range of samples; or ideally, for
more direct comparisons using a wide range of stock
identi®cation techniques on the same samples.
Although, the latter approach is the preferred option
to resolving stock identi®cation problems, only one
study has speci®cally examined this approach (Wald-
man et al., 1997). The paucity of studies utilizing such
an approach is due to a number of reasons including
39
the different expertise of individual scientists conduct-
ing the research, the speci®c stock identi®cation issues
and purposes being addressed, and the logistical costs
in utilizing multiple, complementary techniques in a
single study.
Incorporating different stock identi®cation methods
into a single study often allows apparent discrepancies
implied by each method to be resolved. This is illu-
strated by Kinsey et al. (1994). Electrophoretic data
revealed a single panmictic population of Spanish
sardine, Sardinella aurita, in coastal Florida, however,
morphometric and meristic data indicated regional
patterns. The morphological variations were habitat-
speci®c and diminished as the ®sh got older and
migrated throughout the species range. Leslie and
Grant (1990) found no genetic differences between
samples of angler, Lophius vomerinus, from different
areas along the South African coast, while variation in
meristic and morphometric characters was evident.
Differences in these characters were thought to be
environmentally driven and not related to segregation
of speci®c genes. Safford and Booke (1992) concluded
that the phenotypic variation in Atlantic herring,
Clupea harengus harengus, stocks throughout the
northwest Atlantic was most likely due to environ-
mental conditions, as they found no genetic differ-
entiation. Likewise, Pepin and Carr (1993) found no
evidence for genetic discrimination of Atlantic cod,
Gadus morhua, off Newfoundland, in contrast to their
meristic and morphometric data. Extensive egg and
larval drift throughout the region was considered to
have limited the potential for genetic isolation, despite
variation in environmental regimes that may have
affected phenotypic features of larvae as they settled.
When the results of different stock identi®cation
studies are not consonant, the default management
scenario should be to use a precautionary approach to
ensure resource sustainability and maintenance of
genetic biodiversity. In the above mentioned studies,
such an approach was encompassed where each stock
component was recommended to be managed as a
separate unit. Without the information from multiple
techniques an inaccurate determination of stock struc-
ture is possible, although it really depends on why
stock identi®cation is required in the ®rst place (i.e.,
genetic stocks vs. management units).
Relying on a single approach for stock identi®ca-
tion may enable temporal and spatial patterns in stock
40 G.A. Begg, J.R. Waldman / Fisheries Research 43 (1999) 35±44
structure of a species to be followed, but the accuracy
of the technique remains unknown without the use of
additional con®rmatory evidence (Waldman et al.,
1997). For example, Roby et al. (1991) found con-
gruent results of an east±west separation of capelin
stocks in the Gulf of St. Lawrence using truss mor-
phometric and electrophoretic analyses. The conco-
mitant use of different stock identi®cation approaches
improved their con®dence in the results of the stock
identi®cation. Melvin et al. (1992) observed similar
results using mark-recapture, meristic and morpho-
metric data in the discrimination of American shad,
Alosa sapidissima, stocks in the Bay of Fundy. Inte-
grating the meristic and morphometric variables
enabled them to achieve relatively high rates of clas-
si®cation success in mixed-stock analyses. O'Connell
et al. (1995) found signi®cant genetic heterogeneity
between Atlantic salmon, Salmo salar, stocks through-
out Wales using mtDNA and allozyme variation. They
concluded that alternative DNA approaches (as with
other comparative stock identi®cation techniques)
should be compared with results from other techni-
ques to assess the relative power of stock identi®ca-
tion. Johnson et al. (1994) compared electrophoretic
data of king mackerel, Scomberomorus cavalla, in the
Gulf of Mexico to tagging, catch data and spawning
seasonality to help interpret stock identi®cation pat-
terns.
Few stock identi®cation studies have explicitly exam-
ined a comprehensive range of techniques (Casselman et
al., 1981; Waldman et al., 1988; Grif®ths, 1997; Begg et
al., 1998a), and only one has speci®cally assessed the
results of different stock identi®cation approaches in a
statistically rigorous, comparative, fashion (Waldman et
al., 1997). Casselman et al. (1981) examined the stocks
of lake white®sh, Coregonus clupeaformis, in Lake
Huron using a combination of tag-recapture, life history,
morphometric, meristic, and electrophoretic data. Mul-
tiple stocks were identi®ed, with tagging data providing
the best evidence of discreteness between the stocks.
Other characteristics provided additional information
on the speci®c details of the separations. Mark-recapture
data delineated ®ve stocks, while the life history
parameters indicated that another six groups were
relatively discrete and probably constituted separate
stocks, even though some of the differences may have
been in¯uenced by changes in environmental condi-
tions.
Waldman et al. (1988), in a review of stock identi-
®cation techniques used for striped bass, concluded
that the best overall resolution between stocks
occurred for a combination of approaches that have
high individual resolutions, but low mutual congru-
ence (i.e., techniques that do not have a substantial
degree of parallelism in the type of variation that is
measured). Grif®ths (1997) found con®rmatory evi-
dence from several life history parameters, morpho-
metric relationships, and mark-recapture data used to
identify South African stocks of silver kob, Argyro-
somus inodorus. Begg et al. (1998a) investigated the
stock structure of two mackerels, Scomberomorus
queenslandicus, and, S. munroi, off the Australian
east-coast using tag-recapture, life history, otolith
chemistry, electrophoretic and commercial catch data.
Integrating complementary information from the dif-
ferent approaches was essential in interpreting the
results from any one character, particularly for the
phenotypic-based attributes, where a number of
hypotheses (e.g., environmental differences) could
have been speculated to have caused the spatial (stock)
patterns, rather than simply genetic factors.
Only one empirical comparison of alternative stock
identi®cation approaches ± utilizing the same speci-
mens for each technique has been conducted to date
(Waldman et al., 1997). In that study the morpho-
metric-based or genotypic approach best characterized
mixed-stocks of striped bass. Their results also sug-
gested that estimates of stock composition may be
inaccurate if a phenotypic-based approach does not
have relatively high rates of classi®cation success,
despite the use of correction factors.
A recommended protocol for integrated stock
identi®cation, therefore, is the use of at least a genetic
procedure and at least one phenotypic-based app-
roach. Genetics may be used to provide a direct basis
for stock structuring and to interpret phenotypic-based
patterns (Ihssen et al., 1981; Smith, 1990). As
described above, morphometric and meristic charac-
teristics have been frequently used in association with
different genetic approaches (e.g., Claytor and Mac-
Crimmon, 1987; Lindholm and Maxwell, 1988;
O'Maoleidigh et al., 1988; Riget et al., 1992). The
application of morphological and meristic character-
istics in resolving stock identi®cation problems, how-
ever, is complicated as phenotypic variation may not
be due to genomic differences (Clayton, 1981), but by
 G.A. Begg, J.R. Waldman / Fisheries Research 43 (1999) 35±44
environmental factors (Lindsey, 1964; Todd et al.,
1981). Mark-recapture has also been commonly used
in conjunction with genetic-based studies. Owing to
the logistics and expense of large-scale tagging pro-
jects, these have been usually conducted as separate
studies, or as precursors to genetic analyses (Johnson
et al., 1986, 1994; Ross and Sullivan, 1987). Mark-
recapture data can provide information about ®sh
movements that can assist in de®ning individual
spawning aggregates, which are prerequisites for
genetic-based de®nitions of stock structure. Mark-
recapture data are also useful not only in providing
estimates of movements and stock mixing, but also
growth, mortality and stock size, which are important
parameters for ®shery management (Pawson and Jen-
nings, 1996). Parasite markers have been used in
combination with genetic methods to differentiate
stocks and can provide an indication of stock mixing
similar to mark-recapture techniques (MacKenzie,
1983; Belyaev and Ryabov, 1987; Lester, 1990).
Genetic methods are useful for determining evolu-
tionary history related to stock structure (particularly
for endangered species management), in contrast with
phenotypic markers that are more applicable for
studying short-term environmentally-induced varia-
tion (which are more often used for purposes of ®shery
management) (Roby et al., 1991; Kinsey et al., 1994).
The spatial and temporal scales of management inter-
est de®ne the choice of stock identi®cation techniques.
Genetic-based differences re¯ect evolutionary time
scales and may produce more conservative groupings
(i.e., fewer stocks) than those based on chemical,
morphometric, parasitic or tagging studies (Elliott
et al., 1995).
4. Discussion
Despite the need to reliably identify management
units of the same species, such classi®cation cannot be
accomplished with a single technique (Edmonds et al.,
1989; Campana et al., 1995). Combining the results
obtained with several techniques may provide con-
siderable insight to the possible stock structure of a
species (Elliott et al., 1995). The ability to readily
characterize the identity of an individual ®sh or group
of ®sh taken in a particular area and/or time, to a
particular stock remains a major challenge (Kutkuhn,
41
1981), and will always be probabilistic lacking a
technique or protocol that unambiguously identi®es
the origin of individual ®sh (Waldman et al., 1988). An
holistic approach to ®sh stock identi®cation is highly
desirable owing to the limitations and conditions
associated with any particular method and the require-
ments of ®shery management for separating units
based on genotypic or phenotypic differences. Proce-
dures are now available that can provide a suite of
information on the biology, distribution, and implied
stock structure of a species.
The utility of stock identi®cation techniques should
be considered on a case-by-case basis depending also
on the degree of resolution required. Tagging and
parasitic data generally provide broad-scale stock
identi®cation information, but may be inadequate
for determining more complex multi-stock structures,
unless greater emphasis is placed on obtaining more
thorough recapture information than is typical (MacK-
enzie, 1983; Begg et al., 1997). Morphometrics, mer-
istics and life history characteristics have been used
successfully for stock identi®cation at a range of
different scales (Casselman et al., 1981; Elliott et
al., 1995; Cadrin and Friedland, 1999, these proceed-
ings), but are often limited by their possible alteration
by environmental variation (Lindsey, 1964; Todd et
al., 1981). Chemical methods enable elemental `®n-
gerprints' to be discerned for multiple stock com-
plexes (Campana et al., 1995; Begg et al., 1998b;
Thresher, 1999, these proceedings), although environ-
mental variation may exist within the distribution of a
single genetic stock. Molecular procedures can pro-
vide a genetic basis for stock identi®cation (Smith,
1990; Shaklee et al., 1999, these proceedings). Con-
sequently, the different circumstances related to each
stock identi®cation technique, and the scale at which
stocks can be detected vary depending on the situation.
Application of multiple stock identi®cation techni-
ques (i.e., an holistic approach) to stock identi®cation
problems may con®rm a particular stock structure ®rst
detected by a single procedure used in isolation.
Overlaying all available information from a range
of techniques will enable a generalized, consistent
and de®nitive pattern of stock structure to be devel-
oped, relative to the needs of ®shery management.
Such an approach would enable a higher degree of
con®dence in a particular stock structure, rather than
that for one that had been generated by a single
42 G.A. Begg, J.R. Waldman / Fisheries Research 43 (1999) 35±44
procedure. This would be particularly relevant for
situations where different techniques provide different
interpretations of stock af®nities. In such cases, reli-
ance on a single procedure could provide an inaccurate
representation of the underlying stock structure of a
species, which could have signi®cant management
implications in the utilization and conservation of
its stocks. Importantly, the various means of obtaining
sources of samples and data for alternate stock iden-
ti®cation approaches may often be complementary,
thereby minimizing time and collection costs (Hohn,
1997).
The process of de®ning ®sh stocks is essential for
effective ®sheries management, and will continue to
undergo re®nement as new tools and technologies are
developed (Kutkuhn, 1981). Although, a precise deter-
mination of stock identi®cation remains a major chal-
lenge, and will not necessarily be suf®cient if ®sheries
occur on mixed stocks, our best opportunity at identi-
®cation between these mixed catches relies on exam-
ining all available stock identi®cation information using
the most up-to-date technologies when logistically fea-
sible. The sampling of ®sheries vs. the sampling of ®sh is
a real challenge in linking the results of laboratory-
intensive stock identi®cation techniques to ®sheries
operations (and ®sheries management), and subsequent
impacts on the ®sh stocks. Stock identi®cation should
thus be recognized as a continuing process, evolving as
management needs for stock assessment change, but
always viewed against the background of a critical
examination of all available data and new studies as
dictated by changing resource condition and experi-
mental technologies (Brown et al., 1987).
Acknowledgements
Thanks to Fred Serchuk, Steve Murawski, Kevin
Friedland, Jon Gibson, and William Overholtz for
reviews and critical comments of this paper. This
work was performed while G.A. Begg held a National
Research Council (NOAA/NMFS/NFSC) Research
Associateship.
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