Palaeogeography, Palaeoclimatology, Palaeoecology 514 (2019) 672–676

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Palaeogeography, Palaeoclimatology, Palaeoecology

journal homepage: www.elsevier.com/locate/palaeo

Identifying bias in cold season temperature reconstructions by beetle mutual T

climatic range methods in the Pliocene Canadian High Arctic
⁎
Tamara L. Fletchera,b, , Adam Z. Csankb, Ashley P. Ballantynea
a
Department of Ecosystem and Conservation Sciences, University of Montana, Missoula, MT, USA
b
Department of Geography, University of Nevada, Reno, NV, USA

A R T I C LE I N FO A B S T R A C T

Keywords: Well-preserved beetle elytra from the fossil and subfossil record are used by palaeoclimatologists to estimate past
Palaeoproxy temperatures. Beetle-derived estimates of temperature across the Pliocene Arctic are consistently lower than
Winter those derived from other palaeoclimate proxies. Here we test if that pattern is attributable to either the modern
Vegetation dataset or specific mutual climatic range method (MCRM). We used a global dataset of observations (Global
Coleoptera
Biodiversity Information Facility) for modern beetles as opposed to a North American regional dataset, and a
CRACLE
consistent MCRM for comparison to vegetation-derived estimates, termed CRACLE (Climate Reconstruction
Analysis using Coexistence Likelihood Estimation), rather than different MCRM methods for each analysis.
Beetle-derived estimates from one site, Beaver Pond, are consistently much lower than the vegetation-derived
estimates, stable isotope results from flora and fauna, and summer temperatures from bacterial tetraethers. The
reanalysis of beetle data produced a small reduction in proxy estimate mismatch overall. We found good
agreement among proxy estimates of maximum temperature of the warmest month for two of the three sites in
the Canadian Arctic, but increasingly poorer agreement for warmest quarter, coldest quarter and minimum
temperature of the coldest month. This pattern may be due to beetles' avoidance mechanisms and a climate space
different from modern calibration sets during the Pliocene. Beetle derived temperature estimates, especially cold
season, should be interpreted with caution when reconstructing palaeoclimates.

1. Introduction
All palaeoclimate proxy methods have uncertainties for which ad-
justment must be made, such as taphonomic filtering (Greenwood,
1992; Sangster and Dale, 1961) and phylogenetic constraints (Kennedy
et al., 2014; Steart et al., 2010). The family of palaeoproxies referred to
as Mutual Climatic Range Methods (MCRM) rely on the modern climate
ranges of taxa to reconstruct past climate where a suite of taxa is known
(e.g. Atkinson et al., 1987; Harbert and Nixon, 2015; Mosbrugger and
Utescher, 1997).
One potential problem with applying this method is that the geo-
graphic distribution of the reference taxa used may be incompletely
recorded. This issue was proposed to explain the difference between
climate estimates derived by MCRM for vegetation (Fletcher et al.,
2017; Matthews et al., 2003) and for beetles (Elias and Matthews,
2002) at key sites in the Pliocene Canadian High Arctic (Fig. 1; Gosse
et al., 2017). Despite concerns regarding the suitability of beetles in
coastal Beringia as proxies for the mean temperature of the coldest
month (Tmin-mean; Alfimov and Berman, 2009; Elias, 2010), these
results have nevertheless generally been accepted. Some of the beetles
used to estimate Pliocene High Arctic climate have since been observed
south of their recorded range (J.V. Matthews Jr. pers. obs. 2016; see
Fletcher et al., 2017). Other Beringian taxa have wider thermal ranges
in Eurasia than North America, and thus global datasets are likely more
accurate training data for palaeoclimate estimates.
Another key critique of MCRM is that they are traditionally based on
presence/absence data only, not accounting for abundance. Thus,
MCRM assume a species is equally likely to be found in any part of that
range (Birks et al., 2011; Huppert and Solow, 2004). Thus far, beetle
MCRMs have relied on local calibration to account for changing mi-
crohabitat use across their total range and issues of under- and over-
estimation of temperature at community range boundaries (Alfimov
and Berman, 2009). More recently, abundance-based methods for ap-
plication to beetle assemblages have been developed. For example,
Huppert and Solow (2004) developed a technique that creates summed
probability curves for the climate variable of interest based on quad-
ratic logistical regressions for each species in a database. However,
these probability curves are symmetric and unimodal while the true

⁎
Corresponding author at: Department of Ecosystem and Conservation Sciences, University of Montana, Missoula, MT, USA.
E-mail address: tamara.fletcher@umontana.edu (T.L. Fletcher).

https://doi.org/10.1016/j.palaeo.2018.11.025
Received 2 August 2018; Received in revised form 22 November 2018; Accepted 22 November 2018
Available online 24 November 2018
0031-0182/ © 2018 Published by Elsevier B.V.
T.L. Fletcher et al.
distribution may not follow a Gaussian distribution and could be
strongly skewed.
Bray et al. (2006) suggest the method of Huppert and Solow (2004)
and other MCRMs are biased towards replicating modern climate in-
stead of reconstructing past climates. This is due to over representation
of certain climate spaces in the current climate and thus an increased
probability of observing organisms therein in the modern calibration
set. Bray et al. (2006)'s ‘ubiquity analysis’ uses the frequency that a
species is observed in each quadrant with a matching climatic signal.
This method's drawbacks include subjective criteria for dealing with
marginal data (Bray et al., 2006), although subjectivity was the pro-
blem MCRM was designed to correct (Elias, 2010). Ultimately, Bray
et al. (2006) stated that little could be done in the short-term to produce
the frequency data and distribution range that would improve beetle
MCRM estimates. In the 12 years since Bray et al. (2006), global, multi-
sourced, databases that may provide an approximation of regional
abundance and improved distribution range have been developed that
allow testing of some of these critiques.
In this study, we apply a recently-developed MCRM, CRACLE
(Climate Reconstruction Analysis using Coexistence Likelihood
Estimation; Harbert and Nixon, 2015), that uses observations from an
internationally coordinated, multi-institutional, opensource global
biodiversity database (Global Biodiversity Information Facility; GBIF.
org/what-is-gbif) to reassess palaeoclimate reconstructions in the
Pliocene Arctic based on beetle assemblages. In doing so, we aim to
compare beetle and vegetation proxy temperature reconstructions and
determine the source of the persistent mismatch in estimates.
2. Methods
The MCRM CRACLE derives observation data for each species from
GBIF, and climate data from the global climate database, WorldClim
(http://www.worldclim.org/; Hijmans et al., 2005), for the coordinates
associated with each observation. For each climate variable, the range
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Palaeogeography, Palaeoclimatology, Palaeoecology 514 (2019) 672–676
Fig. 1. A map of the Canadian High Arctic
marking the sample localities. BI – Banks Island
(3.6–5.333 Ma); BP – Beaver Pond (3.9 + 1.5/
−0.5 Ma); MI – Meighen Island (~3.6 Ma; see
Fletcher et al., 2017 for a discussion of age un-
certainties). Map data: Google Earth Pro
7.3.2.5491; International Bathymetric Chart of
the Arctic Ocean, Landsat/Copernicus, and U.S.
Geological Survey.
that is most likely to be inhabited by all taxa in the fossil assemblage is
derived by a joint maximum likelihood approach. This has three ad-
vantages: Firstly, it makes use of global beetle observations acquired
since the publication of the Pliocene Arctic estimates (Elias and
Matthews, 2002); Secondly, the statistical method potentially accounts
for outliers and community boundary effects by including additional
regional abundance information in the modern dataset, thus not re-
quiring local calibration; Thirdly, the spatial resolution of available
climate data has improved: ~4 km compared to 25 km grids. The ve-
getation-derived CRACLE estimates used for comparison here were
previously found to yield empirically similar results to estimates from
isotopic studies, other vegetation-derived methods, and bacterial
branched glycerol dialkyl glycerol tetraethers (brGDGTs; Ballantyne
et al., 2010; Ballantyne et al., 2006; Csank et al., 2011a; Csank et al.,
2011b; Fletcher et al., 2017). The CRACLE code also calculates the
climate interval based on the underlying method, but not database, of
the Coexistence Approach (Mosbrugger and Utescher, 1997; see Tables
S1 and S2 for comparison), instead using the GBIF. CRACLE provides
parametric (P-CRACLE) and non-parametric (N-CRACLE) methods for
deriving climate estimates (parametric results are provided here) in-
cluding mean temperature of the warmest quarter (TwarmQ) and
coldest quarter (TcoldQ), maximum temperature of the warmest month
(Tmax), and minimum temperature of the coldest month (Tmin),
whereas Elias and Matthews (2002) estimates are for the mean tem-
perature of the warmest month (Tmax-mean) and coldest month (Tmin-
mean). The code used in this article is the original code published by
Harbert and Nixon (2015), to be directly comparable with the vegeta-
tion results from Fletcher et al. (2017), though note an updated version
of the code is now available at Github (https://rdrr.io/github/rsh249/
CRACLE/).
Predatory and scavenging beetles were used by Elias and Matthews
(2002) (Table S3) to derive the original MCRM estimates. They were
chosen for their proposed rapid shifts with climate change as compared
with herbivores that are tied to host plant distributions (Elias, 2010),
T.L. Fletcher et al.
Table 1
Summary of the difference in mismatch between the previous (Vegetation CRACLE minus Beetle MCRM) and new (Vegetation CRACLE minus Beetle CRACLE)
estimates for climate using beetles.
Warm season
Vegetation CRACLEa - Vegetation CRACLEa - Vegetation-beetle
beetle MCRM beetle CRACLEb mismatch change
Banks Island 5.3–6.5 °C 0.8–3.5 °C −3 to −4.5 °C
Beaver Pond 5.2–6.3 °C 6.8–8.2 °C +1.6–1.9 °C
Meighen Island 4.4–5.6 °C −0.2–2.3 °C −3.25 to −4.0 °C
a
Average of maximum temperature of the warmest month and mean temperature of the warmest quarter (Tmax and TwamQ) and average of mean temperature of
the coldest quarter and minimum temperature of the coldest month (Tmin and TcoldQ).
b
The average of Tmax and TwarmQ, and Tmin and TcoldQ as the closest equivalent available to the beetle MCRM mean temperature of the warmest month
(Tmax-mean).
and are considered independent of plant proxies (Coope, 2009) that
may be slower to respond. This investigation uses the same predatory
and scavenging taxa as Elias and Matthews (2002) to test the hypothesis
that the mismatch between beetle and vegetation derived temperature
estimates persists even when global observation data is used and the
method is held constant. This is tested by re-estimating the climate
parameters from the beetle taxa using CRACLE (Harbert and Nixon,
2015), and comparing them to vegetation-derived CRACLE estimates
produced by Fletcher et al. (2017).
3. Results and discussion
The application of CRACLE to the Pliocene beetle fauna improved
agreement with the vegetation-derived estimates for maximum tem-
peratures but minimum temperature biases persisted. On average there
was ~6 °C less mismatch between beetle-derived and vegetation-de-
rived CRACLE estimates, than between the estimates from the original
beetle-based MCRM and vegetation-derived CRACLE (Table 1). This
effect was present in the warm season estimates except at Beaver Pond
(BP; 1.6 to 1.9 °C greater mismatch), but was strongest in the cold
season estimates across all sites (mean ~10 °C less mismatch; Fig. 2)
where the difference between beetle and vegetation-derived estimates
were originally much higher (mean ~20 °C). The improved agreement
between estimates may be due to likely increased range data from the
GBIF or increased climate tolerance information using abundance. Al-
ternatively, the Tmin from CRACLE, rather than the Tmin-mean from
the MCRM, may be a better predictor of beetle range as it is the absolute
coldest temperature experienced, even if briefly, that may be lethal
(Alfimov et al., 2003; Elias, 2010).
Despite using the same method and database, the beetle-derived
estimates of temperature for BP were consistently lower than the ve-
getation-derived estimates. Comparing to additional, independent
proxies, brGDGTs, summer temperatures of ~15.4 °C (Fletcher et al.,
2018) are consistent with the vegetation-derived estimate of
14.8–15.7 °C (Fletcher et al., 2017) and not the much cooler 7.5–7.9 °C
TwarmQ derived from the beetle fauna. Similarly, the June–July
growth season temperature estimate from δ18O isotopes in larches
(15.8 ± 5.0 °C; Csank et al., 2011a), and May to September tempera-
ture estimate from isotopes in mollusks (10.2–14.2 °C; Csank et al.,
2011b) are also consistent with the warmer vegetation-derived esti-
mate. We suggest this reflects a skew to cool observations within the
envelope suggested by the coexistence approach (7.0–15.8 °C TwarmQ;
Table S1). To investigate the cause of this skew we conducted a CRACLE
analysis of the BP beetles excluding aquatic types, as recommended by
Alfimov and Berman (2009). The result was a slight increase in tem-
perature (Table 2), more prominent in Tmin and TcoldQ, but in-
sufficient to explain the much cooler temperatures estimated by beetles
compared with other proxies.
For the Meighen Island (MI) and Banks Island (BI) sites, only the
estimates for Tmax overlap in their range and the difference between
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Palaeogeography, Palaeoclimatology, Palaeoecology 514 (2019) 672–676
Cold season
Vegetation CRACLEa - Vegetation CRACLEa - Vegetation-beetle
beetle MCRM beetle CRACLEb mismatch change
17.5–19.6 °C 7.5–12.4 °C −7.2 to −10.4 °C
20.4–21.8 °C 13.2–17.5 °C −4.3 to −7.25 °C
14.7–16.6 °C 2.6–7.8 °C −8.8 to −12.1 °C
climate estimates increases as cooler components of the year are in-
cluded (Fig. 2). As noted by Elias and Matthews (2002), and quantita-
tively shown by Elias et al. (1996), Huppert and Solow (2004) and
others, cold season temperature estimates derived from the beetle fauna
are less constrained than those for the active, warm season period.
Following Harbert and Nixon (2015), we tested beetle-derived CRACLE
against modern sites in North America, calculating the difference be-
tween the CRACLE estimates and WorldClim values. This analysis
confirmed the reduced reliability of beetle-derived CRACLE estimates
for cold season (Tmin and TcoldQ; Table 3) and also showed that this
change in accuracy is most influenced by poor performance at esti-
mating cold season temperatures at the coldest sites (S4). Both beetle
behavioral avoidance strategies (Elias, 2001) and physiology may ex-
plain why they are a poor recorder of cold temperatures. Unlike plants,
the small body size and mobility of most beetles allows them to take
maximal advantage of microhabitat refugia (Danks, 2004), resulting in
observed differences between surface temperature and microhabitat
temperature of at least 40 °C (Baust and Miller, 1970). Rather than
avoiding cold, some insects in extreme regions either control the
structure and location of freezing within the body or facilitate super-
cooling (Lee, 2010) allowing survival in some taxa below −85 °C (Li,
2012, 2016). Due to these adaptations some beetles may not experience
meteorological air temperature during winter or may not be sensitive to
a wide range of temperatures below freezing. The addition of taxa with
wide tolerances, however, should not adversely affect the precision of
the CRACLE method. If included taxa avoid or tolerate a wide range of
cold season temperatures, an increase in the range of estimates for cold
season temperature is expected. Indeed, our CRACLE results for beetles
do reflect a nearly three-fold increase in the estimate range from Tmax
to Tmin. This is expected from the ‘less tight’ relationship between
beetles and Tmin-mean for the MCRM that had been established pre-
viously (Elias et al., 1996), but does not explain a bias to colder esti-
mates.
If the beetle fauna are tightly linked to warm season temperatures
but have a wide range of cold season tolerance, the cold season tem-
perature estimates may reflect the modern seasonal range of tempera-
ture of the calibration dataset rather than the Pliocene cold season. For
example, for sites with similar Tmax in WorldClim to that reconstructed
here by vegetation, the Tmin range is between −52 °C and 14.9 °C, with
a mean of −23.00 °C, skewness of −1.25 and Kurtosis of −0.62 (see
the distribution in Fig. S5). This demonstrates the potential for modern
climate space to bias to cold temperatures for taxa that have a wide
range of winter temperature tolerance. Indeed, (Elias, 1999) found that
the beetle MCRM underestimates Tmin-mean at modern Alaskan
coastal sites compared with observations. A bias to modern continental
observations, which is certainly the case by area, could bias cold season
estimates low; the calibration bias raised by Bray et al. (2006) may not
be overcome by application of CRACLE if the fauna has a wide tolerance
to cold season temperature.
Alternatively, the inclusion of diving aquatic species whose thermal
T.L. Fletcher et al. Palaeogeography, Palaeoclimatology, Palaeoecology 514 (2019) 672–676

Table 2
A comparison of CRACLE temperature estimates derived from beetles at
Pliocene Arctic sites (BI = Banks Island, BP = Beaver Pond, MI = Meighen
Island) from the original taxa list (Elias and Matthews, 2002), with the removal
of aquatic beetles, and with adding phytophagus species, where known in the
palaeofauna.
From original Without Without aquatics but with
taxa list (°C) aquatics (°C) phytophagus species (°C)

Tmax BI 20.5–22.1
BP 13.6–13.9 13.6–14
MI 19.8–21.2 19.9–21.3 20–21.4
TwarmQ BI 12.5–13.8
BP 7.4–7.7 7.4–7.7
MI 12.5–13.6 13.6–14.8 13.8–15
TcoldQ BI −11.3 to −8
BP −21.1 to −18.3 −18.1 to
−15.2
MI −9.4 to −6.3 0–2.9 −0.1–2.9
Tmin BI −16.4 to −13.3
BP −23.5 to −20.4 −20.9 to
−17.8
MI −15 to −11.6 −4.4 to −1.3 −4.4 to −1.4

Table 3
The change in the average minimum difference between the WorldClim value
and CRACLE estimate across modern test sites from the warmest to coldest
periods. Beetle values reported reflect only sites with > 10 taxa, brackets in-
dicate value for all sites. Removing low diversity sites may underrepresent the
coldest sites due to the latitudinal biodiversity gradient (for full details see S4).
Vegetation values from Harbert and Nixon (2015).
Average minimum difference

Beetles Vegetation

Maximum temperature of the warmest month 1.6 °C (2.4 °C) 0.9 °C

Mean temperature of the warmest quarter 1.3 °C (2.3 °C) Not reported
Mean temperature of the coldest quarter 2.6 °C (6.1 °C) Not reported
Minimum temperature of the coldest month 3.7 °C (7.5 °C) 0.7 °C

Fig. 2. The most likely range of the four temperature variables (maximum
temperature of the warmest month, mean temperature of the warmest quarter,
mean temperature of the coldest quarter, and minimum temperature of the
coldest month) as reconstructed from vegetation-derived CRACLE estimates
(leaf columns; light) and beetle-derived CRACLE estimates (beetle columns;
dark).
range reflects deep water temperatures rather than air or surface water
temperatures may bias to colder estimates under some climate condi-
tions, as shown by Alfimov and Berman (2009). To test this we con-
ducted a CRACLE analysis on the MI fauna excluding aquatic species
(no aquatic species were in the BI list). Confirming the effect at BP and
in Alfimov and Berman (2009), the results were warmer than the
temperature estimates with aquatic species included. Indeed, the esti-
mates were warmer than those from the vegetation CRACLE for cold
season, and within range of the warm season estimates – although this
is not universal given the original BI list did not include aquatic taxa.
Finally, Alfimov and Berman (2009) also argued that phytophagus
species should be included in beetle MCRMs. We tested this by adding
the phytophagus species identified at MI by Matthews and Telka
(1997): Chrysomelidae; Donacia distincta, Donacia subtilis, (identified as
Donacia distincta-subtilis group), Curculionidae; c.f. Grypus equiseti
(Fab.), Notaris bimaculatum. Neither BI nor BP have phytophagus taxa
identified to the species level. The effect of adding phytophagus beetles
to the taxa list with no aquatics was less than ± 0.2 °C on either end the
temperature estimate range (Table 2). Overall, the results of different
taxa selection were mixed in these beetle fauna, but removing aquatics
led to some improvement.
A final possible explanation for the mismatch is evolutionary change
in the thermal biology of the beetles. Addo-Bediako et al. (2000) found
that while upper thermal tolerances of insects show little geographic
variation, the lower thermal tolerance shows an increase in variation
with latitude. Elias and Matthews (2002) contend that it is highly un-
likely that the thermal tolerance of the beetles used here has changed,
due to the constancy of species assemblages unlikely to have all evolved
‘in the same direction’. Nevertheless, it is a difficult possibility to either
show or deny.
While high-latitude plants are also adapted to cold, there are rea-
sons to consider their distribution more tightly correlated to winter
temperature. Plants cannot use behavioral strategies to avoid the
coldest temperatures and so rely on freezing resistance, fine-tuning of
plant phenology, and alterations in plant stature – all of which vary by
taxa (Körner, 2016). For example, the timing of the beginning and
length of the growth season is species-specific and plays a critical role in

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T.L. Fletcher et al.
the survival of mature tissue over winter (Körner et al., 2016). Plants
are exposed to winter damage such as desiccation of wind exposed
needles and dormant buds, which varies by species (Havranek and
Tranquillini, 1995). The suite of physiological adaptations to cold vary
in mechanism and effective range between species and even between
tissues of the same species, such that plant climatic ranges may be
meaningfully distinguished below 0 °C (Havranek and Tranquillini,
1995).
Given beetles' adaptations to cold potentially causing wide toler-
ances and the persistent mismatch between other proxies and beetles,
we posit the beetle Tmin-mean estimates from the Pliocene Arctic
should be considered biased towards cold temperatures.
4. Conclusions
The beetle-derived reconstruction of Tmax agrees with other esti-
mates and is likely a robust proxy. The less constrained relationship
between beetles and Tmin-mean than for vegetation, disagreement
between the cold season estimates and those from vegetation, and the
improvement in match between proxies when aquatic species are re-
moved, suggests that the taxa used for the original Pliocene Arctic es-
timations are not good recorders of cold season temperatures during the
Pliocene. Due to differences in seasonality during past global warm
periods, biases in modern calibration data may be amplified, and it is
critical that multiple proxies for climate are used and cautiously eval-
uated.
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.palaeo.2018.11.025.
Acknowledgments
The authors thank Howard Falcon-Lang (editor), Robert Harbert
and two anonymous reviewers for their comments that improved this
manuscript; also a previous editor and two reviewers whose advice
improved an earlier version of this work, and Cameron Wilson for
editing assistance. This work was funded by NSF Division of Polar
Programs (Grant 1418421) to A.P.B. and an Endeavour Research
Fellowship (5928-2017) to T.L.F.
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