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A trait-based approach to thermal niches: linking movement and metabolism

to predict thermal tolerances of terrestrial invertebrates

Preprint · February 2022

DOI: 10.22541/au.164371451.19184810/v1

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 1 A trait-based approach to thermal niches: linking movement and

2 metabolism to predict thermal tolerances of terrestrial invertebrates

3 Jördis F. Terlau1,2, Thomas Boy1,2, Ulrich Brose1,2, Benoit Gauzens1,2, Samraat Pawar3,

4 Malin Pinsky4, Myriam R. Hirt1,2

5 1. EcoNetLab, German Centre for Integrative Biodiversity Research Halle-Jena-Leipzig,

6 Leipzig, Germany

7 2. Institute of Biodiversity, Friedrich‐Schiller‐University Jena, Jena, Germany

8 3. Department of Life Sciences, Imperial College London, Silwood Park, Ascot, UK

9 4. Department of Ecology, Evolution and Natural Resources, Rutgers University, New

10 Brunswick, NJ, USA

11

12 Running title: Movement and metabolism drive thermal niches

13 Keywords: Feeding rates, costs of transport, image-based tracking, encounter rates, trophic

14 interactions, ectotherms, net energy gain

15 Type of article: Letter

16 Number of words in the abstract: 139

17 Number of words in the main text: 4921

18 Number of references: 109

19 Number of figures: 5

20
21 Corresponding author: Jördis F. Terlau, German Centre for Integrative Biodiversity

22 Research (iDiv) Halle-Jena-Leipzig, Puschstraße 4, 04103 Leipzig, Germany, +49 341

23 9733240, joerdis_franziska.terlau@idiv.de

24

25 E-mail addresses of co-authors:

26 Myriam R. Hirt: myriam.hirt@idiv.de

27 Thomas Boy: thomas.boy@posteo.de

28 Ulrich Brose: ulrich.brose@idiv.de

29 Benoit Gauzens: benoit.gauzens@idiv.de

30 Samraat Pawar: s.pawar@imperial.ac.uk

31 Malin Pinsky: malin.pinsky@rutgers.edu

32

33 Statement of authorship:

34 JFT, UB and MRH conceptualized the study. TB gave technical support during the

35 experiments. JFT conducted experiments and analyzed output data. MRH and JFT wrote the

36 first draft of the manuscript and all authors contributed substantially to revision.

37

38 Data accessibility statement: Should the manuscript be accepted, the data will be archived

39 in an appropriate public repository and the DOI will be included at the end of the article.
40 Abstract

41 The survival of animals under global warming strongly depends on their individual thermal

42 niches, which result from the balance between energy loss and gain. Active movement is an

43 important component of this energetic balance, as it affects not only energy gain via food

44 intake but also energy loss via activity metabolism.

45 Here, we develop a novel trait-based approach for how thermal niches arise from

46 temperature-dependent movement. Therefore, we used image-based tracking to quantify the

47 unimodal responses of the movement speed of carabid beetles to temperature. We used these

48 empirical data to parameterize a mathematical model based on metabolic and predator-prey

49 theory for net energy gain to derive a general mechanistic concept of thermal niches. This

50 trait-based approach allows a relatively rapid and cost-effective assessment of climate change

51 vulnerability for a wide range of animal taxa on broad geographic scales.

1
52 Introduction

53 We are globally confronted with climate change, which manifests itself in a rapid rise in

54 average temperature and extreme temperature events (Jones & Mann 2004; IPCC 2021).

55 Temperature is one of the key drivers affecting ecological patterns and processes. This

56 includes direct effects on organismal physiology and performance (Huey & Kingsolver 1989;

57 Angilletta 2009) as well as indirect effects via changes in habitat characteristics and resource

58 availability (Parmesan 2006; Durant et al. 2007). In particular, the strong direct temperature

59 dependence of basic physiological processes, such as metabolism, growth, and reproduction,

60 renders ectotherms most threatened by climate warming (Gillooly et al. 2001; Brown et al.

61 2004; Dillon et al. 2010; Dell et al. 2011; Lemoine & Burkepile 2012). To assess such

62 sensitivity, the temperature tolerance of an organism is typically described as a performance

63 curve with a lower thermal limit, an optimum temperature, and an upper thermal limit,

64 yielding the so-called thermal niche (Huey & Stevenson 1979). The upper and lower limits of

65 these ecological thermal niches are often defined as the temperatures at which population

66 growth rate is null (Colinet et al. 2007; Berger et al. 2008; Deutsch et al. 2008; Angilletta

67 2009). Previous studies have used physiological traits like respiration and aerobic

68 performance or the isotherms that limit the geographic distribution of species to derive

69 thermal niches (Pörtner & Knust 2007; Kearney & Porter 2009; Sánchez-Fernández et al.

70 2016; Verberk et al. 2016; Arostegui et al. 2019). However, it is impossible to empirically

71 measure these traits for the plethora of ectothermic species worldwide. This calls for a

72 mechanistic approach to thermal niches grounded in fundamental trait-based processes that is

73 more generalizable across species.

74 The underlying mechanism driving population growth rates is the balance between energy

75 gain and energy loss. Energy loss is mainly driven by the individual metabolic rate

76 determining the average energy expended over a specific period, which increases

2
 77 exponentially with temperature (Gillooly et al. 2001; Brown et al. 2004; Ehnes et al. 2011).

78 To compensate for this increased energy loss through warming, animals need to enhance their

79 energy gain by higher food intake (Pörtner et al. 2006; Lurgi et al. 2012; Lang et al. 2017).

80 Movement constrains the fundamental ability to access spatially distributed or mobile

81 resources (Bonte & Dahirel 2017; Goossens et al. 2020) and is thus one of the major

82 processes driving trophic interactions (Dell et al. 2014b; Schlägel et al. 2020). Movement in

83 general and speed in particular have been shown to increase with temperature in ectotherms

84 (Angilletta et al. 2007; Gibert et al. 2016; Cloyed et al. 2019) leading to higher encounter and

85 subsequent attack rates (Aljetlawi et al. 2004; Dell et al. 2011; Pawar et al. 2012; Rall et al.

86 2012). Thus, an increase in movement speed with higher temperatures directly increases an

87 organism’s feeding and energy gain. However, movement is also costly and, thus,

88 temperature increases species energy loss not only through an increase in metabolic rate but

89 also through additional costs of elevated movement speed (Halsey 2016).

90 The individual thermal niche can be described by the net energy gain (NEG, energy gain -

91 energy loss) and covers the temperature range at which NEG is positive (Fig. 1). While

92 energy loss via metabolism increases exponentially, individual movement and consumption

93 rates as well as population growth rates often show a left-skewed unimodal shape across

94 prokaryotes and eukaryotes (Smith 2011; García-Carreras et al. 2018; Hurford et al. 2019)

95 (Fig. 1a).

96 However, the shape as well as magnitude of the specific scaling relationships of energy loss

97 and gain have important implications for the resulting thermal niche (Huey & Kingsolver

98 2019) (see Fig. 1b showing two examples of possible relationships of energy gain with

99 temperature).

100

3
101 Here, we develop a mechanistic, trait-based approach to predict movement-based thermal

102 niches from the allometric and thermodynamic dependence of energy gain and loss. As we

103 are still lacking a general temperature response of movement speed across a wider body-mass

104 range (Dell et al. 2011), we first measured the movement speed of differently-sized carabid

105 beetles under varying temperature regimes using automated image-based tracking. We then

106 integrated this empirically evaluated response of movement speed to temperature into the

107 calculation of feeding rates and metabolic costs across a temperature gradient. This allowed

108 us to mechanistically derive how thermal niches of ectotherms arise from temperature-

109 dependent movement and how this ultimately affects their survival under warming on a broad

110 geographic scale.

111

112 Materials and Methods

113 Study organisms and experiment

114 To quantify the temperature-dependence of movement speed, we measured the movement of

115 carabid beetles (Carabidae) in the laboratory using automated image-based tracking (Dell et

116 al. 2014a; Barnes et al. 2015). We used the following species of carabid beetles: Carabus

117 nemoralis, Carabus granulatus, Pterostichus cristatus, Pterostichus melanarius, Abax

118 parallelus, Nebria brevicollis, Harpalus affinis, and Anchomenus dorsalis. For a detailed

119 description of collection as well as keeping of the beetles, the experimental setup and the

120 tracking process, see supplementary material section (1).

121 Before starting a film recording session, we weighed each individual and kept the beetles

122 separately in small boxes with perforated lids. Followed by an acclimation time of two hours

123 in the environmental chamber, we released one single beetle into the arena. After a time delay

124 of ten minutes to account for the temporarily open doors of the climate chamber, a one-hour

125 film recording was initiated.

4
126 We used two levels of relative humidity (50% and 70%) across a temperature gradient of 14

127 levels from 8 °C to 32 °C. During the recording, we kept a constant temperature and took

128 three separate records for every temperature level using different individuals. In total, we

129 recorded movement, weight, and temperature data for 125 individuals.

130

131 Analyses and statistics

132 We assume a power-law scaling of speed [mm s-1] with body mass (Hirt et al. 2017a, b) and

133 compare two models for the response to temperature [°C]:

134

135 𝑛𝑜𝑛 − 𝑢𝑛𝑖𝑚𝑜𝑑𝑎𝑙 𝑟𝑒𝑠𝑝𝑜𝑛𝑠𝑒 𝑡𝑜 𝑡𝑒𝑚𝑝𝑒𝑟𝑎𝑡𝑢𝑟𝑒: 𝑣𝑇 = 𝑒 𝑎 𝑀𝑏 𝑒 𝑐𝑇 (1a)

2 /𝑤
136 𝑢𝑛𝑖𝑚𝑜𝑑𝑎𝑙 𝑟𝑒𝑠𝑝𝑜𝑛𝑠𝑒 𝑡𝑜 𝑡𝑒𝑚𝑝𝑒𝑟𝑎𝑡𝑢𝑟𝑒: 𝑣𝑇 = 𝑒 𝑎 𝑀𝑏 𝑒 −(𝑇−𝑧0 −𝑧) (1b)

137

138 where e is the base of the natural logarithm, 𝑎 is the intercept (in log space), 𝑀 is body mass

139 [mg], 𝑏 is the allometric exponent, 𝑇 is temperature [°C], and c is the temperature response

140 coefficient for a species (modeled as a random effect) in the non-unimodal model. For the

141 unimodal model, we additionally allowed for species-specific temperature optima with 𝑧0 as

142 the group's average optimal temperature, 𝑧 as the optimal temperature deviation for each

143 species from the group mean (modeled as a random effect), and 𝑤 as the width of the thermal

144 response curve. After log-transformation, we fitted a mixed effects model to the data by

145 applying the nlme function of the nlme package (Pinheiro et al. 2021) with ln(mass),

146 temperature, width of the thermal response curve and 𝑧0 as fixed effects and 𝑧 as a random

147 effect. Note that all following predictions were based on the species average optimal

148 temperature. Thus, 𝑧 = 0 for our calculations of energy gain and loss. We compared the two

149 models using the Bayesian Information Criterion (BIC).

150

5
151 Calculation of net energy gain and thermal niche

152 To illustrate the effects of temperature-dependent movement on energy gain, loss, and the

153 thermal niche of terrestrial invertebrates, we expanded our calculations to represent a

154 predator-prey pair. We used the parameters for handling time from Vucic-Pestic et al. (2011),

155 particularly the data on three different carabid beetles feeding on flightless adults of

156 Drosophila hydei. Thus, in this exemplary calculation, the predator and prey have body

157 masses of MC = 111.75 mg (mean body mass of all beetles used in our study) and MR = 1.91

158 mg, respectively, yielding a body-mass ratio of 58.51. We first used these values for

159 illustrating our concept and then carried out a sensitivity analysis across other body-mass

160 ratios.

161

162 Energy loss

163 To calculate energy loss via metabolism, we used field metabolic rate 𝐼𝑓 [Jh-1], which is the

164 metabolic rate while moving. It comprises the basal metabolic rate and the net costs of

165 transport 𝐶𝑂𝑇𝑛𝑒𝑡 [Jm-1] multiplied by speed 𝑣𝐶 [m h-1] (Peters 1993):

166 𝐼𝑓 = 𝐶𝑂𝑇𝑛𝑒𝑡 𝑣𝐶 + 1.2 𝐼𝑏 (2)

167 We calculated the net costs of transport using 𝐶𝑂𝑇𝑛𝑒𝑡 = 11.3 𝑀𝐶 0.72 (after Peters 1993) with

168 consumer body mass 𝑀𝐶 [kg]. The factor of 1.2 multiplied by the basal metabolic rate 𝐼𝑏

169 accounted for additional costs for lifting the organism’s body from the ground (see Peters

170 1993). The predator speed, 𝑣𝐶 , followed equation (1a) or (1b) depending on our empirical

171 movement results (see results for equation and parameter values). To calculate the basal

172 metabolic rate 𝐼𝑏 [J h-1], we used a subset for carabid beetles of the data of Ehnes et al.

173 (2011), which yielded the following equation:

𝑇−𝑇0
𝐸𝑥
174 𝐼𝑏 = 𝑖𝑜 𝑀𝐶𝑏 𝑒 𝑘𝑇𝑇𝑜 (3)

6
175 with 𝑙𝑛 (𝑖0 ) = −5.94, 𝑏 = 0.74 and 𝐸𝑥 = 0.63. Here 𝑖0 is a normalization constant, body

176 mass 𝑀𝐶 [mg], 𝑏 is the allometric exponent, 𝐸𝑥 [eV] is the rate’s activation energy, 𝑇 is the

177 temperature [K], 𝑘 [eV K-1] is Boltzmann’s constant (8.63 x 10-5 eVK-1) and 𝑇0 [K] the

178 normalization temperature (here, 0 °C, i.e., 273.15 K).

179 Assuming that an organism has periods of activity and inactivity, we included an activity

180 factor (𝛼) into our calculations of total energy expenditure: 𝐼𝑡𝑜𝑡𝑎𝑙 = 𝛼 𝐼𝑓 + (1 − 𝛼) 𝐼𝑏

181 As an example, we chose an 𝛼-value of 0.5, implying 50% activity. The total metabolic

182 expenditure, thus, results from 50% activity metabolism and 50% basal metabolism at rest.

183 Since actual activity periods may differ, we also performed a sensitivity analysis on the

184 effects of activity on upper and lower thermal limits (see Supplementary Figure S1).

185

186 Energy gain

187 To calculate the energy gain via feeding, we used per capita feeding rates, 𝐹 [Ind. h-1],

188 depending on prey density, 𝑁 [Ind. m-2] (Holling 1959):

𝑎𝑁
189 𝐹= (4)
1+𝑎ℎ𝑁

190 Here 𝑎 [m² h-1] is the attack rate after Pawar et al. (2012) depending on dimensionality of

191 interaction (here 2D), relative movement speed 𝑣𝑟 [m h-1] of predator (𝑣𝐶 ) and prey (𝑣𝑅 ) and

192 detection distance 𝑑 [m]:

193 𝑎 = 2𝑣𝑟 𝑑 (5)

194 𝑣𝑟 = √𝑣𝑅2 + 𝑣𝐶2 (6)

195 The detection distance was calculated based on the supplementary material of Pawar et al.

196 (2012), giving the equation

197 𝑑 = 100.21 𝑀𝐶 0.36 (7)

7
198 with 𝑀𝐶 as predator body mass [kg]. We calculated the speeds of predator as well as prey, 𝑣𝐶

199 and 𝑣𝑅 , using equation (1a) or (1b) depending on our empirical movement results (see results

200 for equation and parameter values).

201 Additionally, we multiplied the attack rate by a unitless success rate, 𝑠, after Li et al. (2017)

202 to account for an attack success:

𝑅 𝜆
𝑅 1−
𝑅𝑜𝑝𝑡
203 𝑠 = 𝑏0 (𝑅 𝑒 ) (8)
𝑜𝑝𝑡

204

𝑀
205 Where 𝑏0 is the allometric scaling, 𝑅 is the predator-prey body-mass ratio (𝑀𝐶 ) and its
𝑅

206 optimum 𝑅𝑜𝑝𝑡 = 100 and a shaping parameter, 𝜆 = 1. Here, 𝑏0 = 0.1.

207 The handling time ℎ [h Ind-1] follows the equation after Vucic-Pestic et al. (2011):
𝑇−𝑇0
𝐸ℎ
208 ℎ = ℎ0 𝑀𝑏ℎ 𝑒 𝑘𝑇𝑇0 (9)

209 Here, the intercept is at temperature 𝑇0 of 17.5 °C (290.65 K) corresponding to the mean of

210 the temperature gradient that was used in Vucic-Pestic et al. (2011) ℎ0 is a constant, 𝑏ℎ is the

211 allometric exponent, 𝐸ℎ the activation energy [eV], 𝑘 Boltzmann’s constant, and 𝑀𝐶 is body

212 mass [mg]. From Vucic-Pestic et al. (2011) we obtained the following fixed parameters:

213 𝑙𝑛 (ℎ0 ) = 2.99, 𝑏ℎ = −0.66, 𝐸ℎ = −0.24.

214 We predicted the prey density 𝑁 [Ind. m-2] based on the prey body mass:

215 𝑁 = 𝑒 𝑛 𝑀𝑅𝑏 (10)

216 where 𝑛 is a constant, 𝑀𝑅 is prey body mass [mg] and 𝑏 is the allometric exponent. Fitting

217 equation (10) to data of Ehnes et al. (2014) yielded the following parameter values: 𝑛 =

218 −0.65, 𝑏 = −0.71.

8
219 To express the final energy gain, we multiplied the per capita feeding rates [Ind. h-1] by prey

220 body mass 𝑀𝑅 [mg] to yield per capita biomass feeding rates [mg h-1]. We then converted kg

221 wet mass of prey to energy (after Peters 1993): 1𝑘𝑔 𝑤𝑒𝑡 𝑚𝑎𝑠𝑠 = 7 ∗ 106 𝐽.

222 This yields a per capita energy feeding rate F [J h-1]. Finally, we multiplied the feeding rate

223 by the unitless predator’s assimilation efficiency, 𝜀, recalculated for carnivores after Lang et

224 al. (2017):

𝑏
225 𝜀 = 𝑒 έ0 𝑀𝐶 𝜀 (11)

226 Here έ0 is a normalization constant, 𝑀 is body mass [g] and 𝑏𝜀 the allometric exponent.

227 Following Lang et al. (2017), we used the following parameter values: έ0 = −0.12 and 𝑏𝜀 =

228 0.0096.

229 Similar to our calculations of total metabolic expenditure, we also included an activity factor

𝑎𝑁
230 (𝛼) for an exemplary activity of 0.5 in our calculations of energy gain: 𝐹𝑡𝑜𝑡𝑎𝑙 = 𝛼 (1+𝑎ℎ𝑁).

231 Therefore, the total energy gain results from a 50% feeding rate.

232 Finally, we calculated the net energy gain by subtracting the field metabolic rate (energy loss)

233 from the feeding rate (energy gain):

234 𝑁𝐸𝐺 = 𝐹 ∗ 𝜀 − 𝐼𝑓 . (12)

235 To include upper and lower temperature ranges in our predictions for energy scaling and

236 NEG, we used the nls function in R to fit equ. 1b and the predictNLS function from the

237 propagate package(Spiess 2018) to predict the mean as well as the confidence intervals

238 (CI) with a 95% confidence level.

239 As energy requirements and feeding rates are highly dependent on resource availability

240 (Durant et al. 2007; Sentis et al. 2012; Hooten et al. 2019) as well as the body-size ratio of

241 predator and prey (Brose et al. 2008; Vucic-Pestic et al. 2011; Rall et al. 2012) we

242 additionally analyzed how the scaling of NEG with temperature changes across gradients in

243 prey density and body-mass ratios.

9
244 We then used this exemplary calculation for a carabid beetle to predict its thermal

245 performance under current and future climatic conditions. We focused on areas of

246 occurrences of those species, which we used in this study (GBIF.org 2021), i.e. the palearctic

247 and nearctic regions. To define the species distributional range, we used the R package

248 rangemap (Cobos et al. 2021) applying the concave hull method. We used the data of

249 climatologies at high resolution for the Earth land surface areas (CHELSA, Karger et al.

250 2017). To calculate NEG under current temperature conditions (𝑁𝐸𝐺𝑐 ), we used data on the

251 mean temperature of the warmest quarter (years 1997-2013) (Karger et al. 2018). To

252 calculate NEG under future temperature predictions (𝑁𝐸𝐺𝑓 ), we used the predicted mean

253 temperature of the warmest quarter for years 2060-2080 with a moderate greenhouse gas

254 emissions scenario, Representative Concentration Pathway (RCP) 4.5 (Karger et al. 2020).

255 Following Knutti et al. (2013) and Sanderson et al. (2015), we selected the following five

256 prediction models: CESM1-BGC, GISS-E2H, IPSL-CM51-MR, MIROC5 and MPI-ESM-

257 MR. These models differ substantially in terms of code and assumptions and therefore

258 capture a range of structural model uncertainty.

259 To include global variation in prey density, we predicted prey density depending on

260 temperature (Antunes et al. 2021, dryad repository). We then calculated the respective

261 present NEG and the future ΔNEG as 𝛥𝑁𝐸𝐺 = 𝑁𝐸𝐺𝑓 − 𝑁𝐸𝐺𝑐 , (see Supplementary Fig. S3-

262 S7). For visualization purposes, we depicted the mean of all models predicting ΔNEG (Fig.

263 5), but see the Supplementary Figures S3-S7 for individual results.

264 All statistical analyses and calculations were performed using R 4.0.3 (R Core Team 2020).

265

10
266 Results

267 Temperature response of exploratory speed

268 To investigate how average exploratory speed responds to temperature, we analyzed tracking

269 data of 125 carabid beetles, belonging to eight species. Body masses ranged from 10 mg

270 (Anchomenus dorsalis) to 303 mg (Carabus granulatus). Comparison between the unimodal

271 and non-unimodal models for the response of speed to temperature (equ. 1a, b) indicated that

272 the former is more appropriate (equ. 1b BICp = 171.86 vs 1a BICnp = 233.36) (Fig. 2).

273 Therefore, we used the unimodal temperature-speed model (equ. 1b) with 𝑣𝑇 =

274 𝑒 3.72 𝑀0.097 𝑒 −(𝑇−24.63 − 𝑧/180.37) and z = 0 for all subsequent analyses (see Table S3 for

275 species-specific z values).

276

277 Net energy gain and thermal niche

278 To understand how temperature-dependent movement affects the thermal niche of

279 ectotherms, we used our empirical results on movement speed (Fig. 2) to calibrate a

280 mathematical model (equ. 1-12) that predicts species-specific thermal response of net energy

281 gain. In this model, the unimodal relationship of movement speed with temperature affects

282 both energy gain (through feeding interactions) and loss (costs associated with movement).

283 Figure 3 shows the resulting responses of energy gain and loss to temperature. The feeding

284 rate also follows a unimodal relationship with temperature (Fig. 3a, energy gain). The field

285 metabolic rate (FMR) depends on the basal metabolic rate and the costs of transport, which

286 increase proportionally to movement speed. Consequently, higher movement speeds with

287 increasing temperature initially raise metabolic costs. At higher temperatures, however,

288 movement speed decreases again and FMR and energy loss are then mainly driven by the

289 exponential temperature-dependence of the basal metabolic rate (Fig. 3a, energy loss). Thus,

290 both energy gain and loss initially increase with temperature, while energy gain via feeding

11
291 exceeds energy loss via metabolism. Feeding rates, however, decrease again at higher

292 temperatures, finally leading to an intersection with the scaling of FMR, which marks the

293 point of starvation and the upper thermal limit (Fig. 3a). Calculating the net energy gain

294 (NEG, energy gain – energy loss, equ. 12) yields a unimodal and left-skewed thermal niche,

295 which first increases to an optimum at 17.36 °C (-0.35/+0.58 °C, 95% CI) and then decreases

296 steeply (Fig. 3b). The upper thermal limit is reached at a temperature of 31.53 °C (-

297 1.49/+1.65°C, 95% CI) for our exemplary predator (body mass = 111.75 mg, with a predator-

298 prey body-mass ratio of 58.51), and represents the point where compensation of energy loss

299 via metabolism by energy gain via feeding becomes insufficient. In this exemplary

300 calculation, we used an activity factor (α) of 0.5 indicating an activity period of 50%. Our

301 sensitivity analysis shows that lower and upper thermal limits are shifted to higher

302 temperatures by increased activity, and the total net energy gain increases with higher α-

303 values (Supplementary Fig. S1).

304 This example (Fig. 3) applies to a fixed prey density calculated from equ. 10. We additionally

305 calculated NEG not only depending on temperature but also on prey density and analyzed

306 their interactive effects.

307 Across temperatures, we observed an increase in NEG with increasing prey density, which

308 levels out towards an asymptote (Fig. 4a). The initial increase in NEG steepens with higher

309 temperatures (-10 to 30 °C), suggesting enhanced energetic gains at low prey densities in

310 warm environments. Moreover, the total NEG increases with temperature, thus making more

311 energy available to be invested in growth and other processes. However, at very high

312 temperatures (35 to 40 °C), the initial increase in NEG with prey density reverts to being less

313 steep and the prey density required for survival (NEG > 0) also increases abruptly, suggesting

314 that temperature extremes must also be compensated for by higher prey densities.

12
315 Moreover, the breadth of the thermal niche also increases with higher prey densities and the

316 upper thermal limit is shifted towards higher temperatures (Fig. 4b). Similarly, maximum

317 NEG, the upper thermal limit, and the width of the thermal niche increase with predator-prey

318 body-mass ratios up to a certain ratio. At higher ratios, however, maximum NEG and

319 subsequently also the upper thermal limit decrease again. At the highest body-mass ratios

320 (here a body-mass ratio of 200), there is no survival possible (Fig. 4c). This implies an

321 energetically optimal body-mass ratio of predator and prey (here a ratio of 32 with a

322 maximum NEG at 21.5°C), while too small or too big resource items ultimately result in

323 insufficient feeding rates to compensate for metabolic energy losses at high temperatures.

324 The three-dimensional surface plot (Fig. 4d) shows the interactive effects of temperature and

325 prey density (similar results for body-mass ratios are not shown). On the one hand, higher

326 prey densities provide higher chances of survival and may therefore shift the upper thermal

327 limit towards higher temperatures. On the other hand, higher temperatures increase NEG up

328 to an optimum, followed by a rapid decrease.

329 To project the effects of future climate warming, we calculated the current NEG and the

330 change in NEG (ΔNEG , future NEG – current NEG) for a predator body mass of 111.75 mg

331 and a body-mass ratio of 58.51 under future conditions (climate projection for the years

332 2060-2080, Fig. 5). For geographic mapping, we used the areas of occurrence of the species

333 in this study (see Supplementary Fig. S2 for occurrence data). Our results show that the

334 consequences of temperature increases are neutral for some regions (ΔNEG of approx. zero,

335 white regions), and beneficial for others (positive ΔNEG, blue color scale). An increase in

336 NEG means that with increasing temperature the organisms gain energy in [Jh-1] compared to

337 current conditions. For many regions, however, temperature increases are detrimental

338 (negative ΔNEG, red color scale). This happens when future NEG is lower than current NEG,

339 implying that energy expenditure is still increasing while energy gain remains constant or is

13
340 already decreasing. Regions with a negative ΔNEG are more vulnerable to temperature

341 increase and occur especially in the northern mid-latitudinal regions (e.g. Mediterranean

342 region).

343

344 Discussion

345 To enable trait-based generalizations of thermal niches across species, we developed an

346 empirically-informed mathematical model based on animal movement, which drives energy

347 gain by enabling encounters with resources (Pawar et al. 2012; Dell et al. 2014b) and energy

348 loss via the costs associated with locomotion (Peters 1993; Alexander 2005).

349 Empirically, we showed that the average exploratory speed of carabid beetles follows a

350 unimodal relationship with temperature. We used this relationship to parameterize our model

351 and derive the thermal niches of species depending on their body mass by defining the

352 temperatures at which gains outweigh losses. Our modeling results show that energy loss and

353 energy gain both initially increase with temperature with feeding rates exceeding energy loss

354 via metabolism. Feeding rates, however, decrease again at higher temperatures, resulting in

355 less energy gain than loss and potential starvation at a specific thermal limit. Empirical

356 measures of feeding and attack rates have also shown a unimodal, temperature-dependent

357 relationship (Englund et al. 2011; Vucic-Pestic et al. 2011; Rall et al. 2012; Sentis et al.

358 2012; Betini et al. 2019). In our model, the unimodal relationship of feeding rates arises from

359 the empirically derived unimodal response of movement speed and thus encounter rates with

360 temperature. By simultaneously accounting for the thermal response of basal and field

361 metabolism, the model provides a mechanistic derivation of the commonly observed left-

362 skewed thermal performance curves of individual and population growth rates (Frazier et al.

363 2006; Kingsolver & Woods 2016), and the thermal response of fitness (Amarasekare &

364 Savage 2012).

14
365 There are multiple factors influencing the shape and width of these thermal niches. For

366 instance, the availability of resources alters feeding rates and thus energy gain (Avgar et al.

367 2011; Rall et al. 2012; Calcagno et al. 2014). We have shown that with increasing prey

368 density, (1) the total net energy gain increases independent of temperature and (2) the thermal

369 optimum as well as the upper thermal limit of the thermal niche are shifted towards higher

370 temperatures. The width of the thermal niche also increases concurrently. Similarly, the

371 interactive effects of temperature and prey density (Fig. 4d) show that the effects on net

372 energy gain can be either beneficial or detrimental, depending on the baseline temperature

373 and resource availability. This implies that animals living in areas with a high prey density

374 have a higher capacity to cope with temperature increases. Moreover, they are more resilient

375 to temperature fluctuations due to the increasing width of the thermal niche. This parallels

376 findings of models and experiments showing that reduced food intake lowers optimum

377 temperatures and upper thermal limits (Brett 1970; Thomas et al. 2017; Huey & Kingsolver

378 2019; Huxley et al. 2021). In light of the predicted reduction in resource availability with

379 global change (Ciais et al. 2005; Vasseur & McCann 2005; Zhao & Running 2010;

380 Blanchard et al. 2012; Lister & Garcia 2018), ectotherms will thus become less heat tolerant

381 while simultaneously experiencing higher environmental temperatures (Huey & Kingsolver

382 2019). Eutrophication, however, might counteract this negative effect. For example, animals

383 have been shown to migrate towards the poles in response to global warming (Parmesan

384 1996, 2006; Parmesan et al. 1999; Hughes 2000). In this vein, eutrophication might enable

385 terrestrial animals to migrate faster or maintain a positive population growth for longer in

386 their native habitats. Contrary, warming and eutrophication have both been shown to decrease

387 oxygen availability in aquatic environments, which lowers the upper thermal limit (Pörtner &

388 Knust 2007; Deutsch et al. 2015). Extensions of our model approach to marine environments

389 should thus also account for oxygen availability.

15
390

391 We have also illustrated how our approach can be used to project changes in net energy gain

392 under current and future climatic conditions. A decrease in net energy gain implies that

393 ectotherms have less energy available for reproduction and growth and a higher risk of

394 crossing their upper thermal limits, which jointly reduces their overall chances of survival. In

395 contrast, critical or lethal temperatures, for which immobility or mortality are often measured

396 at extremely high temperatures, already exceed an ecological relevant range (Terblanche et

397 al. 2011; Bennett et al. 2018, 2021). Our approach, however, applies an ecologically relevant

398 and applicable measurement of thermal limits. Our predictions of net energy gain and thus

399 the probability of species occurrence under current climatic conditions match the distribution

400 range of the species from our study quite well. However, our model overestimates the

401 distribution range particularly towards higher latitudes. More precise predictions would

402 require additional information on, for example, habitat requirements, eco-evolutionary

403 processes, and data on local prey availability. However, this simplified gain-loss model

404 already provides a reasonable estimate of potential distribution ranges based only on body

405 mass.

406 Under future conditions, our model predicts strong decreases in net energy gain in general,

407 particularly in the southern parts of the distribution area. In the northern parts, however,

408 animals may even experience a slight increase in net energy gain. This result matches other

409 studies suggesting variable impacts of temperature increases on thermal performance with

410 tropical and Mediterranean organisms being particularly vulnerable to future temperature

411 increases (Vasseur & McCann 2005; Deutsch et al. 2008; Kingsolver et al. 2013; Sinclair et

412 al. 2016; Payne & Smith 2017). A major reason for the higher susceptibility of tropical and

413 mediterranean species to climate warming is related to the exponential increase of metabolic

414 rates with temperature (Dillon et al. 2010; Huey & Kingsolver 2019). Species from cold

16
415 regions experience a smaller increase in metabolic rate for the same increase in temperature

416 and thus suffer less energy loss than species from warm regions. This also matches recent

417 findings showing reduced persistence of species in warm regions compared to species in

418 colder temperate and frigid zones (Gauzens et al. 2020) as well as global patterns of thermal

419 extinction rates of ectotherms due to lowered population growth rates caused by changes in

420 timing of activity (Sinervo et al. 2010). However, predictions of extinction risks that are

421 based on metabolism alone are simplistic, because these organismal loss rates can be

422 balanced or reversed by positive temperature effects on energy gain through feeding rates.

423 Our movement-based approach quantitatively predicts changes in metabolism as well as

424 feeding rates with temperature, and thus offers a flexible tool to relate such patterns to first-

425 principle processes in a relatively tractable and cost-effective way.

426

427 Our predictions of thermal niches are currently based on metabolic rates and movement data

428 of central European carabids. Therefore, the resulting net energy gains and thermal niches do

429 not account for variance across taxonomic, geographic, and evolutionary backgrounds.

430 However, the unimodal response of movement to temperature is likely a general

431 pattern across species and realms (Kamykowski & McCollum 1986; Deere & Chown 2006;

432 Dell et al. 2011; Martin et al. 2012; Monaco et al. 2017; Rezende & Bozinovic 2019;

433 Cecchetto et al. 2020). Thus, our approach could be applied to other taxonomic groups and

434 geographic areas by appropriately modifying the model parameterizations.

435 Further extensions of our approach could also consider resource quality (Ilić et al. 2021),

436 which has been shown to increase feeding rates with decreasing resource quality as

437 compensatory feeding (Ott et al. 2012; Jochum et al. 2017) and therefore impact the net

438 energy gain. Additionally, there are indications that warming will alter feeding behavior like

439 the selectivity for resources (Lemoine & Burkepile 2012; Sentis et al. 2014; Gauzens et al.

17
440 2021), and thus organisms may choose energetically preferable food to compensate for

441 increased energy loss due to warming (Gordon et al. 2018). A promising direction for future

442 research will be to combine effects of changes in resource stoichiometry on assimilation

443 efficiencies (Jochum et al. 2017; Manzoni et al. 2017).

444 Furthermore, our model predictions on future net energy gain so far neglect potential

445 mechanisms for adapting to temperature change. This includes changes in behavior like the

446 use of thermal refuges (e.g. shade in vegetation or habitat structure) (Kearney et al. 2009),

447 shifts in activity times (Dynesius & Jansson 2000; Davis & Shaw 2001; Gunderson & Leal

448 2016) as well as reductions in metabolic rates or body size in response to warming (Gardner

449 et al. 2011; Marshall & McQuaid 2011). Nonetheless, our concept provides a novel approach

450 to assess thermal niches based on movement, which can be flexibly modified and expanded

451 by various aspects and research directions.

452

453 Conclusion

454 The sheer abundance and diversity of invertebrates reveal their functional importance on

455 Earth (Wilson 1987). In order to understand and predict the future impacts of warming on

456 invertebrates, it is essential to gain a mechanistic understanding of how basal temperature-

457 dependent processes affect their temperature tolerance. Here we have leveraged the fact that

458 movement is a key process that underpins animal life in the real world and simultaneously

459 drives the temperature-response of potential food intake and energy expenditure, to quantify

460 the thermal niche of ectotherms. Geographic mapping of the predictions of these thermal

461 niches suggests that global warming may favor species at high latitudes while putting their

462 low latitude populations at risk. This mechanistic approach links our understanding of

463 thermal niches to trait-specific movement processes and thereby provides an integrative

18
464 framework for generalized predictions of thermal niches for the plethora of invertebrate

465 species across different climates.

466

467 Acknowledgements

468 J.F. Terlau, T. Boy, U. Brose, B. Gauzens, M. Pinsky, and M.R. Hirt acknowledge the

469 support of the German Centre for integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig

470 funded by the German Research Foundation (DFG, FZT 118). J.F. Terlau was supported by

471 the research group DynaCom (spatial community ecology in highly dynamic landscapes:

472 from island biogeography to metaecosystems) funded by the German Research Foundation

473 (DFG, FOR 2716). M. Pinsky acknowledges support from NSF #DEB-1616821.

474

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750

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751 Figure captions

752

753 Figure 1: Figure 1: Conceptual figure illustrating how temperature-dependent movement

754 affects an organism’s energetics. Temperature on the x-axis and required energy and net energy

755 gain (NEG) on the y-axis. (a) shows the scaling of energy loss and gain. The grey solid line

756 represents energy loss. The black solid line shows a unimodal increase in energy gain with

757 temperature (A). The black dashed line shows an exponential increase of energy gain with

758 temperature (B). The hatched blue area represents the respective net energy gain. (b) shows

759 that the resulting thermal niche is either a left-skewed thermal niche where NEG first increases

760 slowly towards an optimum followed by a rapid decrease at high temperatures (A), or a bell-

761 shaped niche with a relatively low magnitude of NEG across the thermal range (B). The dashed

762 vertical red and blue lines represent upper and lower thermal (physiological) limits

763 respectively.

764

32
765 Figure 2: The unimodal response of exploratory speed [mm s-1] to temperature [°C] of

766 carabid beetles is best predicted by a unimodal model (equ. 1b, a = 3.72 ± 0.18 [mean ± SE],

767 b = 0.097 ± 0.04, w = 180.37 ± 23.39, z0 = 24.63 ± 1.29; see Table S3 for the species-specific

768 optimal temperature deviation z from the group mean). Colored curves show

769 species-specific optimal temperature response curves with the respective optimal

770 temperatures indicated by triangles at the bottom. The black curve and triangle show the

771 temperature response and optimal temperature average across species, respectively

772

33
773 Figure 3: (a) The thermal response of energy use of an exemplary predator based on

774 temperature-dependent movement (equ. 1b). The solid purple line indicates the energy loss

775 (field metabolic rate, equ. 2-3) and the orange line energy gain (feeding rate, equ. 4-11) and

776 the solid purple line the energy loss (field metabolic rate). (b) Resulting thermal niche based

777 on net energy gain (energy gain – energy loss). The grey line shows the upper thermal limit

778 of an exemplary predator with 111.75 mg body mass feeding on a prey of 1.91 mg body

779 mass. Shaded areas show the 95% confidence intervals. For visualization purposes, we set all

780 negative NEGs (i.e., where energy loss exceeds energy gain) to zero.

781

34
782 Figure 4: (a) The dependence of net energy gain (NEG) on prey density at different

783 temperatures (color code yellow to brown). (b) The dependence of NEG on temperature for

784 different prey densities (color code yellow to dark blue). (c) The dependence of NEG on

785 temperature for different predator-prey body-mass ratios [C/R] (color code light pink to dark

786 pink). Filled points in (b) and (c) show the respective upper thermal limits. (d) Two views of

787 a three-dimensional surface plot showing the interactive effect of prey density and

788 temperature on NEG.

35
789 Figure 5: (a) Map of net energy gain (NEG in [Jh-1]) of a temperate carabid beetle (M ~ 111

790 mg) under current climate conditions. (b) Map of change in NEG for years 2060-2080 as

791 compared to the present (1997-2013). Calculations are based on the mean temperature of the

792 warmest quarter predicted from five different climate models (see Supplementary Figures S3-

793 S7). In panel (b), blue areas indicate an increase in NEG; red areas indicate a decrease in

794 NEG. Dashed borders in (a) and (b) demarcate areas of occurrences of the species, which

795 were used in this study. These occurrences are based on GBIF data and processed using a

796 concave hull type.

797

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