Edge thermal niche tracking
We estimated edge thermal niche tracking by testing whether the change
in minimum or maximum temperature at the range edge over time was
different from zero, based on 90% Bayesian credible intervals from
Bayesian linear regressions of temperature on time. Of 153 species range
edges, we found that 131 (86%) maintained at least one component of the
edge thermal niche (warm or cold extreme temperature) during the study
period. Further, for the majority of range edges—111, or 73%—both
minimum and maximum temperatures were maintained over time. Of the 20
range edges consistent with only one temperature metric, 13 were
consistent with cold extremes and 7 were consistent with warm extremes.
In the West Coast and the Eastern Bering Sea, almost all range edges
tracked both temperature metrics (84% and 97%, respectively; Figure
3B, 3C). By contrast, only 34% of range edges in the Northeast tracked
both temperature metrics, and the Northeast also contained 20 of the 22
range edges that did not track either warm or cold temperature extremes
(Figure 3A). Lack of tracking arose for different reasons in poleward
and equatorward edges: the poleward edges that did not maintain their
edge thermal niches often did not shift as the oceans warmed, or even
shifted south, while the equatorward edges that did not maintain their
edge thermal niches often shifted north faster than expected, into
cooler waters (Appendix 4).
DISCUSSION
We quantified temporal dynamics and thermal niche tracking over decades
for more than 150 marine range edges while using a novel spatiotemporal
modeling approach to standardize among the three study regions and their
differing survey methodologies. Across diverse geographies, historical
climates, and taxa, range edges of marine species were in general
conserving their thermal niches through space and time. Edge thermal
niche conservatism suggests that most species range edges are tracking
temperature change, consistent with evidence that many species
distributions are shifting through space (Pinsky et al. 2013, Hiddink et
al. 2015, Day et al. 2018, Lenoir et al. 2020) and supported by theory
from thermal ecophysiology (Sunday et al. 2012, Stuart-Smith et al.
2017, Pinsky et al. 2019). However, a non-negligible number of range
edges did not shift as predicted—especially in the Northeast, the
region with the greatest historical temperature increase—indicating
that temperature alone does not explain range edge dynamics for all
marine species. This study provides the first large-scale, multi-region
analysis of thermal niche tracking at range edges and describes novel
statistical approaches that are applicable to a wide range of taxa and
systems.
Species ranges are shifting poleward around the globe, both on land and
in the sea (Chen et al. 2011, Poloczanska et al. 2013). Local patterns
in climate change have helped to explain distributional shifts of many
species, especially those that did not shift directly toward the poles
(Pinsky et al. 2013, Lenoir and Svenning 2015). These findings suggest
that species’ range shifts can be at least partially explained by
spatial shifts in their climatic niches (Loarie et al. 2009, Burrows et
al. 2011). While many global change studies have not measured range edge
displacement, those that have often report major poleward shifts in
range edges, particularly at the poleward range edge (Parmesan et al.
1999, Hickling et al. 2005). However, most studies on range edges have
used a small number of time points (e.g., Hickling et al. 2005)—and
often just two—limiting inference about climatic niche tracking.
Marine species are predicted to track their climatic niches more closely
than terrestrial species: they have exhibited greater range shifts to
date, and are more physiologically vulnerable to warming (Pinsky et al.
2019, 2020, Lenoir et al. 2020). Testing for climatic niche tracking is
fundamentally related to measuring range edge dynamics,
because—especially in the oceans—range edges are expected to
coincide with climatic niche limits, specifically thermal limits (Sunday
et al. 2012).
By measuring thermal niche conservatism at the range edge, we tested for
a relationship between range edge position and the isotherms
representing winter and summer temperatures over time (Sunday et al.
2015, Fredston‐Hermann et al. 2020). A finding that a range edge
remained in the same winter or summer temperatures over time can have
several interpretations. It does not necessarily imply that the isotherm
moved: an edge may track its thermal niche either by shifting in the
same direction and at the same rate as an isotherm, or by remaining in
place when the isotherm is stationary. The two regions in our study with
high levels of edge thermal niche tracking, the West Coast and the
Eastern Bering Sea, both had relatively little temperature change when
averaged over the study period; thus, range edges in those regions that
did not move much were typically classified as tracking the edge thermal
niche.
Recent work on marine heatwaves has underscored the need to move beyond
means to measure climatic extremes and variability in studies of global
change biology, including in the oceans (Smale et al. 2019). We
quantified edge position in relation to temperature extremes precisely
for this reason, especially given the marked recent increases in warm
extreme (summer) temperatures in the Northeast and the Eastern Bering
Sea. Yet our results revealed that range edges remained in the same cold
extreme (winter) temperatures approximately as often as they did for
summer temperatures, suggesting that winter temperatures may be an
underappreciated covariate of range dynamics at both poleward and
equatorward range edges (Dana 1853, Morley et al. 2017). Foundational
biogeographic theory provides a hypothesis for this: poleward range
edges could be influenced either by summer temperatures limiting
reproduction and growth or by winter temperatures limiting survival; and
equatorward range edges could be influenced either by summer
temperatures limiting survival or by winter temperatures limiting growth
and reproduction (Hutchins 1947). Further work could test whether this
is a biologically plausible explanation for these temperate marine
species. Longer time series, extensive analysis of different dimensions
of temperature change, or additional oceanographic data products such as
high-resolution hindcast sea bottom temperature data could be used in
the future to tease apart more precisely which temperature metrics best
explain range edge dynamics and why.
Our finding that the region with the greatest historical temperature
increase exhibited the lowest frequency of edge thermal niche tracking
underscores the critical importance of considering non-temperature and
indirect processes that may influence species distributions. In the
Northeast, we documented equatorward range edges that shifted much
further north than expected based on temperature—into cooler
waters—and poleward range edges that did not shift or shifted south
(Appendices 3 and 4). This could arise due to density-dependent habitat
selection if these species were declining in abundance, causing each
range edge to collapse toward the range center (Blanchard et al. 2005).
At the equatorward edge, competition or predation from the south could
be driving edge retraction (Kordas et al. 2011); for example, the rapid
contraction of the equatorward range edge of American lobster could be
due to increased predation from species like black sea bass shifting up
the coast (McMahan and Grabowski 2019) or increased mortality from a
temperature-related disease (Groner et al. 2018). At the poleward edge,
species interactions (HilleRisLambers et al. 2013), priority effects
(Fukami 2015), dispersal limitation (Poloczanska et al. 2011), or a lack
of non-thermal habitat (McHenry et al. 2019) could all inhibit northward
shifts. Sessile invertebrates are particularly vulnerable to dispersal
limitation if prevailing currents do not align with local climate
velocities, as in the Northeast (Molinos et al. 2017, Fuchs et al.
2020). Changes in non-temperature abiotic drivers such as dissolved
oxygen are likely to also influence range edge dynamics (Deutsch et al.
2015, Howard et al. 2020).
This study is the first to use a spatiotemporal modeling approach to
estimate range edge dynamics and estimate a standard error around range
edge positions, which we see as important methodological advances. Using
the VAST model, we calculated rates of range edge shift that were
similar in magnitude to those calculated from raw survey data in the
Northeast (Fredston‐Hermann et al. 2020). Our results are not directly
comparable to previous work, however, because—unlike analyses of raw
distribution data—VAST attributes some variation in recorded
observations and abundances to observation error. Continued extensions
of VAST and similar models to distribution data will facilitate more
rigorous evaluation of historical range edge dynamics, even for datasets
with known inconsistencies and biases in sampling.
While we found that range edge positions almost always maintained their
edge thermal niche, year-over-year temperatures at the range edge were
often variable (Appendix 4), and near-term (i.e., annual, not
multi-decadal) projections of a species’ edge thermal niche are unlikely
to predict exactly where it will shift. Further, a non-negligible
fraction of range edges did not shift at all, shifted opposite the
predicted direction, or “overshot” temperature change and shifted into
cooler waters. Our methods provide a blueprint for assessing whether
range edges have tracked their thermal niches and for identifying
species of concern that do not appear to be shifting as expected based
on temperature alone. Classifying species by edge thermal niche tracking
can inform management and conservation because different interventions
are likely required for a species that shifts in response to warming
(e.g., transboundary management) than for a species that remains
stationary in the face of warming (e.g., assisted migration). To move
beyond categorizing all results of no thermal niche tracking as
“individualistic responses,” future research can test edge thermal
niche conservatism against—or jointly with—other biogeographic
hypotheses that integrate the influence of species interactions,
population dynamics, eco-evolutionary processes, and other important
abiotic and biotic drivers. Future progress on range edge dynamics will
be accelerated by mechanistically testing predictions about which
temperature and non-temperature processes should be limiting for which
range edges against biogeographical data. Testing multiple
contemporaneous processes (including density-dependent range expansion
and contraction) and their net effects will provide insight into, and
ultimately enable prediction of, range edge dynamics in a changing
climate.
ACKNOWLEDGMENTS
We thank current and past NOAA staff for collecting, curating, and
publishing the data used in this study. E. M. Howard and R. Mendelssohn
provided expert advice regarding oceanographic and spatial data
analysis. We also thank A. J. Allyn and D. Ovando for code review and
feedback.
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FIGURES