Reports

Ecology, 91(9), 2010, pp. 2499–2505

Ó 2010 by the Ecological Society of America

Breaking Bergmann’s rule: truncation of

Northwest Atlantic marine fish body sizes
JONATHAN A. D. FISHER,1,2,3 KENNETH T. FRANK,2 AND WILLIAM C. LEGGETT1
1
Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6 Canada
2
Ocean Sciences Division, Bedford Institute of Oceanography, P.O. Box 1006, Dartmouth, Nova Scotia B2Y 4A2 Canada

Abstract. A strictly species-centric view of human impacts on ecological communities may

conceal important trait changes key to ecosystem functioning and stability. Analyses of body
size and community composition data for 326 Northwest Atlantic fish species sampled across
.900 000 km2 over three decades revealed a rapid and widespread reduction of body sizes
driven by declines within species and changes in relative abundances. The changes were
unrelated to species richness but of sufficient magnitude to eliminate biogeographic scale
gradients of increasing body size with latitude commonly characterized as Bergmann’s rule.
These changes have persisted despite reduced potential for intraspecific competition and
favorable bottom water temperatures, both of which should lead to increased growth rates.
The aggregate body sizes in these Northwest Atlantic fish communities may now represent a
mismatch between the environmental variability characteristic of the Northwest Atlantic and
the historical body size, life history traits, and productivity of species across this region. We
discuss how these changes may jeopardize the potential for recovery of these important
temperate/subarctic ecosystems.
Key words: Bergmann’s rule; body size; macroecology; marine fish; Northwest Atlantic; temporal
dynamics.

INTRODUCTION Angilletta et al. 2004). Harvesting and environmental

Body size is a key functional trait both within and
among species that influences population abundance,
geographic distribution, species interactions, and physi-
ological performance (Peters 1983, Pauly 1998, Hildrew et
al. 2007). It has been extensively studied from a
biogeographic perspective and forms the cornerstone of
one of the most commonly documented macroecological
patterns, Bergmann’s rule: the general trend for animal
sizes to increase with latitude (Blackburn et al. 1999).
While the original definition related to endotherms
(Bergmann 1847, Watt et al. 2010), many factors have
been identified as potential contributors to the mainte-
nance of spatial gradients in body size in both endotherms
and ectotherms (e.g., Lindsey 1966, Millien et al. 2006,
Olson et al. 2009), and evolutionary adaptations to
latitudinal variability in the habitats occupied are likely
to play a central role (Leggett and Carscadden 1978,
Manuscript received 15 October 2009; revised 5 March 2010;
accepted 24 March 2010. Corresponding Editor: D. E.
Schindler.
3
E-mail: jonathan.fisher@queensu.ca
2499
change can also rapidly modify evolved body size patterns
within species and assemblages (Bianchi et al. 2000, Shin
et al. 2005, Millien et al. 2006, Darimont et al. 2009).
However, it remains unclear whether such changes have
the potential to disrupt the adaptive pattern in body sizes
reflected in Bergmann’s rule by degrading contrasts in
body size across latitudes.
Within the ocean, the world’s largest biogeographic
region, body size plays a fundamental functional role in
structuring trophic interactions (Hildrew et al. 2007). On
a worldwide scale, marine fisheries are largely size-
selective, targeting and/or incidentally removing the
largest individuals and species (Bianchi et al. 2000, Shin
et al. 2005), often due to minimum size requirements for
harvest (Fenberg and Roy 2008). In these exploited
ecosystems the potential exists for a contraction of body
size distributions that may lead to functional homogeni-
zation, an increasing overlap in the functionality of
species that were formerly discrete (Olden et al. 2004,
Jennings et al. 2008).
Here we document the functional elimination of an
historically strong gradient of increasing body sizes with
latitude across three Northwest Atlantic large marine
 2500 JONATHAN A. D. FISHER ET AL. Ecology, Vol. 91, No. 9

FIG. 1. Dynamics of aggregate species geometric mean mass among four-year time blocks across three Northwest Atlantic large
Reports

marine ecosystems illustrating the breakdown of the gradient between body size and latitude (Bergmann’s rule) and the
homogenization of sizes across ecosystems. Offshore boundaries of three Northwest Atlantic large marine ecosystems show the
latitudinal sampling extent from North Carolina, USA (358 N), to Newfoundland and Labrador, Canada (558 N).

ecosystems (LMEs), each .160 000 km2 in area (Ap-
pendix A) within a sampled list of .300 species. This
decline in and homogenization of body sizes occurred
rapidly (within decades) and involved both targeted and
nontargeted species. As a consequence, species that were
once functionally discrete, due to differences in body
size, may now be experiencing greatly intensified
interspecific competition. This may explain why, in spite
of recent dramatic reductions in commercial exploita-
tion rates and improving environmental conditions,
average fish body sizes in the ecosystems involved have
remained depressed. The current body size attributes of
species occupying these ecosystems suggest a mismatch
between the environmental variability characteristic of
these LMEs and the evolved body sizes, life history
traits, and historical productivity of species occupying
these important temperate/subarctic ecosystems.
METHODS
Spatial and temporal variation in Northwest Atlantic
Continental Shelf, Scotian Shelf, and Newfoundland–
Labrador Shelf ) (Fig. 1). For these analyses, the LME
boundaries were defined by the extent of the samples
within the survey database (Fisher et al. 2008). Data
from latitudes 358 N to 558 N and depths and areas
described previously (Fisher et al. 2008) were binned
into seven four-year groups (1975–2002). Binning
provided a temporally integrated portrayal of size
differences that included a period (1991–1994) of
northern groundfish fisheries collapses (Leggett and
Frank 2008). The 1995–2002 Newfoundland–Labrador
Shelf data were collected using different survey methods
(McCallum and Walsh 1997), another reason for
binning by four-year groups. Within each 18 latitudinal
interval b and each four-year time block t, an average
fish biomass m was determined for each sample (total
biomass divided by total number of individuals). The
geometric mean, GM, was then determined as
!
1X n

fish size were examined using an existing database of
marine fish species identities, abundances, and masses
recorded from 27 224 samples collected by fisheries-
independent trawl surveys over three decades (Fisher et
al. 2008). The subset of years (1975–2002) examined
included a total of 24 658 samples (Appendix A; see
Plate 1). Samples were collected by randomly trawling
within predetermined areas stratified by depth within
three adjacent large marine ecosystems (Northeast U.S.
ðGMÞb;t ¼ exp lnðmi Þb;t ð1Þ
n i¼1
where n is the number of samples in latitude b and time
block t. This calculation provided biomass-weighted
geometric means. However, because other studies
investigating latitudinal changes in body size did not
weight species by biomass (e.g., Lindsey 1966, Mac-
pherson and Duarte 1994, Pauly 1998), we also
estimated a biomass-independent geometric mean for
September 2010 NORTHWEST ATLANTIC MARINE FISH SIZES
comparative purposes (Appendix B). The number of
species occurring within each four-year block and 18
latitude was also calculated based on a total of 326 fish
species, but excluded 36 relatively rare species (contrib-
uting 1.2% of total individuals) due to lack of mass data
for these species in all years and regions (Appendix C).
The annual mean masses (total mass/total abundance)
of a subset of 120 species, present within the sampled
region in all seven four-year groups, were examined at the
LME spatial scale to test (t tests) for intraspecific
differences in body mass between years within the first
and second halves of the survey time series. To determine
whether changes in aggregate body masses were a
reflection of relative species abundance and/or species
composition over the 28-year time period investigated, we
examined both standardized anomalies of abundance
data using principal components analysis (PCA; Choi et
al. 2005) and Bray-Curtis percentage similarities in
species composition based on presence/absence data
(Clarke and Warwick 2002) scaled relative to the
composition of the 1975–1978 time block. Together, these
analyses reveal the inter- and intraspecific contributions
to the dynamics of aggregate fish mass.
RESULTS DISCUSSION
AND
The aggregated mean body size of fish species sampled
exhibited a progressive, .10-fold increase from low to
high latitudes throughout the 1970s and mid-1980s (Fig.
1). This pattern is consistent with Bergmann’s rule and
with reports for other groupings of marine (Lindsey
1966, Macpherson and Duarte 1994) and terrestrial
species (Millien et al. 2006, Olson et al. 2009, Watt et al.
2010). However, in the early 1980s the latitudinal
pattern of increasing body sizes began to erode and by
1991, the gradient in mean fish sizes was eliminated and
mean body sizes had declined dramatically (Fig. 1;
Appendix D). This homogenization of aggregate body
sizes across latitudes represents a breakdown of
Bergmann’s rule and was similarly reflected in alternate
portrayals of the data (Appendix B). At the northern-
most latitudes (Newfoundland–Labrador Shelf ), where
the ecosystem effects of overfishing were most severe
(Frank et al. 2006) and where community-wide changes
have been previously reported (Haedrich and Barnes
1997), these changes occurred before the 1995 gear
modification in the Newfoundland–Labrador region (see
Methods) and have persisted in the other two regions
where no gear changes occurred (Appendix A).
In contrast to the strong temporal variation in body
sizes across latitudes, latitudinal variation in species
richness (Fig. 2; Appendix D) and species composition
(based on presence/absence data; Appendix D) remained
relatively stable through time. However, progressive and
reciprocal changes did occur in the relative abundances
of large- and small-bodied species in the areas and over
the time period studied (Fig. 3). This interspecific change
in relative abundances contributed partly to the
reduction in aggregate body size. For example, the
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FIG. 2. Dynamics of cumulative species richness among
four-year time blocks and degrees latitude across three
Northwest Atlantic large marine ecosystems illustrating relative
temporal stability in species richness in comparison to body size
dynamics shown in Fig. 1.
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proportion of samples lacking large-bodied individuals
(.1 kg) rose dramatically in the two northerly LMEs
(Appendix A) and collapses in the biomasses of large-
bodied species that occurred in the early 1990s (average
reduction of 80% in six northern areas; Frank et al.
2006) were followed by orders of magnitude increases in
the biomasses of smaller-bodied (and formerly prey)
species (Choi et al. 2004, Petrie et al. 2009).
Intraspecific declines in body mass were also profound,
and no one geographic area was immune to reductions in
body size. Approximately half of the species present
within the sampled region in all seven four-year groups (n
¼ 61; from 13 orders) showed differences (P , 0.10) in
body mass between time periods within at least one LME
(Appendix E); at this P value ;19 tests may be expected
to exhibit false positives. Within each LME the frequent-
ly occurring species generally declined in body size (mean
change 17%, range 99% to þ221%, n ¼ 192). The
further subset that showed significant changes (mean
change 22%, n ¼ 119; Fig. 4) together comprised 46–
95% of the biomass in the three LMEs (Appendix F). The
magnitude of the change in body size of exploited species
(Atlantic cod, pollock, other gadiform species; Appendix
E) was consistent with published reports of body size
reductions derived independently from ageing data
(Leggett and Frank 2008, Shackell et al. 2010). However,
these declines in mean body size were not confined to
targeted taxa (Fig. 4). Rather, of the 61 species that
showed significant changes (33 commercially targeted
and 28 nontargeted species), most exhibited significant
declines in body size (Fig. 4; Appendix E), which may be
a consequence of the nonselective nature of mobile gear
fisheries within these LMEs. Together, both changes in
interspecific composition within LMEs (Fig. 3) and
 2502 JONATHAN A. D. FISHER ET AL. Ecology, Vol. 91, No. 9

declining trends were evident (Leggett and Frank 2008).

This suggests that in addition to fishery effects,
environmental influences, notably declining water tem-
peratures during the late 1980s/early 1990s, contributed
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FIG. 3. Principal components analysis of annual species-

specific numbers per tow within three large marine ecosystems
(LMEs). (Top panel) The first principal component (PC1)
through time based on standardized anomalies (percentage
values indicate explained variation) of fish abundances. In all
three LMEs, changes in the sign of PC1 occurred in the late
1980s or early 1990s, consistent with changes in body sizes (Fig.
1). (Bottom panel) Box plots based on the maximum total fish
length (data available online at hhttp://www.fishbase.orgi) for
those species that had either strong positive (r  0.5) or strong
negative (r  0.5) loadings on PC1. Positive and negative
correlations indicate increasing and decreasing abundance
through time, respectively. Numbers above box plots indicate
the number of species within each box plot. Open circles show
all species lengths, whiskers indicate the 10th and 90th
percentiles, box boundaries represent the 25th and 75th
percentiles around the median, and solid circles indicate data
outliers.

intraspecific declines in the sizes of dominant species (Fig.

4) influenced the aggregate patterns.
The reductions in mean body sizes of the commer-
cially targeted species cannot be explained by changes in
their individual (population-level) abundances. Virtually
all experienced dramatic declines in abundance and even
collapses during the 1990s and beyond (Haedrich and
Barnes 1997, Petrie et al. 2009) and, notwithstanding
moratoria on fishing imposed in 1992/1993, remain at
very low population densities (Shelton et al. 2006).
Growth data for the species investigated are limited, but
for four species for which reliable size-at-age data exist,
FIG. 4. Magnitude of change in body size among 61 fish
species, including both targeted (circles) and nontargeted species
(squares). Percentage change is based on the difference in annual
mean masses between years within the first and second halves
(1975–1988 vs. 1989–2002) of the sample period except that the
Newfoundland–Labrador Shelf included only 1989–1994 in the
second half. Solid symbols represent significant (P  0.10, chosen
to increase power to detect differences) changes in body size.
Results are presented for (a) the Newfoundland–Labrador Shelf
(18 species significant), (b) the Scotian Shelf (25 species
significant), and (c) the Northeast U.S. Continental Shelf (41
species significant). Changes in mass specific to species and to
large marine ecosystems are provided in Appendix E.
September 2010 NORTHWEST ATLANTIC MARINE FISH SIZES
PLATE 1. A single trawl survey sample from the Scotian Shelf, illustrating the multi-species catch and wide range of marine fish
body sizes. Photo credit: K. T. Frank.
to the decreases in mean sizes within species (which has
been tested simultaneously with density-dependent
growth limitations; Swain et al. 2003). Additionally,
temperature-mediated changes in community composi-
tion have been described in other exploited large marine
ecosystems. For example, compositional changes and a
shift toward dominance by larger sized species were
associated with a warm-water regime in the Gulf of
Alaska (Anderson and Piatt 1999). However, given the
current low densities and warmer-than-average bottom
water temperatures in the Northwest Atlantic that have
now prevailed for more than a decade (Kelly et al. 2009),
evidence of increased growth rates and mean body size
should now be evident. No such increases in growth or
mean body size have been observed (Leggett and Frank
2008, Shackell et al. 2010).
One hypothesis for the contemporary persistence of
small body sizes across the three LMEs is the operation
and intensification of density dependence at the assem-
blage level. This hypothesis is consistent with the
negative relationship between size and local abundance
exhibited by the smaller-bodied species (Appendix G)
and the failure of the once-large-bodied species to
achieve increased growth and body sizes at reduced
densities (Appendix E). The once-dominant predators
have, by virtue of their reduced mean body sizes, become
but one component of a more-or-less homogeneous
species complex now locked in an intensified competi-
tion for resources. Such a scenario has been suggested
for North Sea species, where with increasing fishing
mortality, energy demands of different species peak at
younger, overlapping ages (Jennings et al. 2008).
Support for this hypothesis is also found in numerous
reports of the poor physiological condition (an indirect
2503
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measure of foraging success) of species occupying these
Northwest Atlantic LMEs (Choi et al. 2004, Shackell
and Frank 2007, Wheeler et al. 2009), which have
persisted despite improved temperature and feeding
conditions (Kelly et al. 2009, Petrie et al. 2009, Shackell
et al. 2010). The apparently anomalous failure of the
larger-bodied species to recover and reoccupy their
former niches may also relate to a predator–prey role
reversal in which the young life stages of the once-
dominant large-bodied predators have become the prey
of the now numerically dominant smaller-bodied species
(Barkai and McQuaid 1988, Walters and Kitchell 2001,
Petrie et al. 2009). Another possibility is that the small
body sizes of the once-dominant predators reflect the
evolutionary effects of fishing that have been demon-
strated in experiments conducted at relatively small
geographic scales and suggested in some empirical
studies (Jørgensen et al. 2007, Conover et al. 2009,
Darimont et al. 2009). However, the absence of data on
growth rates for more than a few species and reliable
estimates of natural mortalities precludes rigorous
evaluation of alternate hypotheses.
How, and for how long, the existing assemblages of
species and the homogenized size distribution across
species will persist in these characteristically variable and
harsh temperate and subarctic environments is an open
question. Consistent with the predictions of evolutionary
theory (Leggett and Carscadden 1978, Fenberg and Roy
2008), over millennia, these highly variable, northern
environments selected for iteroparity, large body sizes,
and the diverse suite of body sizes and life history
strategies required to buffer populations against frequent
recruitment failure. The contemporary declines in body
sizes we document represent a profound disruption of a
 2504 JONATHAN A. D. FISHER ET AL.
macroecological pattern (Fisher et al. 2010). These once
remarkably productive fishery resources are now largely
unresponsive to conventional management (e.g., Leggett
and Frank 2008) and now share body size characteristics
of assemblages found in less environmentally variable and
less energetically efficient tropical ecosystems that are
characterized by small body size, low biomass, and high
turnover (Lindsey 1966, Macpherson and Duarte 1994,
Pauly 1998). In these Northwest Atlantic temperate and
subarctic ecosystems the breakdown of Bergmann’s rule is
likely to lead to enhanced population, community, and
ecosystem instability.
In the short term, recruitment failures may become
more common (Leggett et al. 2004, Venturelli et al.
2009), local extinction rates may increase, and opportu-
nistic and possibly invasive species may become
abundant or even dominant. In the longer term, some
or all of these ecosystems may regain their former
characteristic diversity of body sizes and ecological roles
if exploitation rates are reduced (Worm et al. 2009).
However, the timelines required for ‘‘phenotypic resto-
ration’’ in wild populations remain unknown due to
complications involving interspecific interactions (Dari-
mont et al. 2009). The prognosis for restoration of
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historical patterns of body sizes in these marine
ecosystems is further limited by the sustained emphasis
of current fisheries management on minimum size limits
(Fenberg and Roy 2008). And, if the persistent small
body sizes of the once-dominant large-bodied predators
are a reflection of fisheries-induced evolutionary chang-
es, recent experimental evidence suggests that many
generations, in the absence of exploitation, will be
required for recovery of their historical body sizes
(Conover et al. 2009). Given these complexities, our
results suggest that the current focus on maintaining or
increasing taxonomic richness per se in biogeographic-
scale analyses of exploited ecosystems (Worm et al.
2006) is unwarranted. We found species richness to be
insensitive as an indicator of ecosystem stability/health
(Fig. 2) and instead suggest a more immediate need for a
focus on restoring and maintaining key species func-
tional traits (Bianchi et al. 2000, Shin et al. 2005,
Venturelli et al. 2009, Worm et al. 2009).
ACKNOWLEDGMENTS
We thank B. Branton, M. Fogarty, D. Kulka, and J. Palmer
for facilitating data exchange or providing data. B. Petrie, N. L.
Shackell, and the reviewers provided many helpful comments.
Funding for this research was provided by NSERC Discovery
grants to K. T. Frank and W. C. Leggett and the Canadian
Department of Fisheries and Oceans.
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APPENDIX A
Temporal changes in size-based fish characteristics and sampling within three large marine ecosystems (Ecological Archives
E091-182-A1).

APPENDIX B
Alternative mass estimate and body mass dynamics (Ecological Archives E091-182-A2).

APPENDIX C
Species lists (Ecological Archives E091-182-A3).

APPENDIX D
Regressions of latitudinal variation in body size and species richness and temporal changes in species composition (Ecological
Archives E091-182-A4).

APPENDIX E
Intraspecific changes in fish mass in 61 species (Ecological Archives E091-182-A5).

APPENDIX F
Contributions of 61 species to biomass within three large marine ecosystems (Ecological Archives E091-182-A6).

APPENDIX G
Annual mean body mass and densities from Scotian Shelf species (Ecological Archives E091-182-A7).