Mahaut Andréaz Dupoué
email: andreaz.dupoue@gmail.com

Victoire Sorlin

Murielle Richard

Jean-François François Le Galliard

Olivier Lourdais

Jean Clobert

Fabien Aubret

Mahaut Sorlin

Mother-offspring conflict for water and its mitigation in the oviparous form of the bimodal lizard Zootoca vivipara

Keywords: Altitude, mother-offspring conflicts, dehydration, ectotherm, parity mode, reproduction Altitude, mother-offspring conflicts, dehydration, ectotherm, parity mode, 43 reproduction Study species, populations and husbandry

de niveau recherche, publiés ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

The European common lizard, Zootoca vivipara, is a small (adult snout-vent length SVL ~ 50 145 -75 mm), widespread lacertid typically found in cold humid peat bogs and heathland habitats 146 from Western Europe to Scandinavia and Eastern Russia [START_REF] Heulin | Further evidence of the existence of 565 oviparous populations of Lacerta (Zootoca) vivipara in the NW of the Balkan Peninsula. 566[END_REF]. Between the 147 15 th and 31 st of May 2018, we caught a total of 134 adult females (105 gravid and 29 non-148 reproductive) from 6 populations distributed throughout the Pyrenees mountain range (Table 149 S1). The reproductive strategy of the Western oviparous form substantially changes with 150 increasing SVL across altitude (Fig. S2), and earlier and multiple clutches as well as earlier 151 age at maturation observed in warmer habitats at lowland elevations (< 300 m) compared to 152 single reproductive event at higher altitudes [START_REF] Heulin | Demography of a bimodal reproductive 568 species of lizard (Lacerta vivipara): survival and density characteristics of oviparous 569 populations[END_REF]. In our study, the altitudinal 153 range (990-1580 m) was strong enough to examine altitudinal variation of reproductive 154 performance (see below), while narrow enough to avoid strong differences in life history 155 strategies.

156

Females were captured by hand and then transferred to the laboratory where they were 157 housed in individual terraria (18 x 12 x 12 cm) with sterilized soil, a shelter, and basking heat 158 until parturition. Each individual was provided a 20-35°C thermal gradient for 6 hours per day 159 (09:00-12:00 and 14:00-17:00) using a 25 W incandescent light bulb placed over one end of 160 each terrarium. They had ad libitum access to water in a petri dish and we further provided 161 water 3 times per day at 09:00, 13:00, and 17:00. We further provided them with 3 162 mealworms (Tenebrio molitor) every two days. Females were kept in these standard 163 conditions until oviposition except during the water restriction experiment (see below 164 experimental design).

165

We characterized environmental conditions for each population, with altitude (Table 166 S1), presence of permanent versus periodic water sources and air temperature, because these 167 measures were likely to shape local adaptations in the regulation of the water balance as 168 documented previously in the viviparous form (Dupoué et al., 2017a). We recorded air [START_REF] Rodríguez-Díaz | Effects of incubation temperature on 632 hatchling phenotypes in an oviparous lizard with prolonged egg retention: are the two main 633 hypotheses on the evolution of viviparity compatible?[END_REF]Foucart et al., 2018). We used three incubators Females were weighed (BM, ± 1 mg) every three days throughout the water restriction period 204 to assess effects of water restriction on short-term changes since BM is an indicator of 205 hydration state [START_REF] Lillywhite | Dehydration and drinking responses 586 in a pelagic sea snake[END_REF]. However, for simplicity, we only examined BM 206 changes (ΔBM = BM final -BM initial ) from the initiation to the end of the water restriction 207 period. We also measured absolute changes in plasma osmolality (ΔOsmo = Osmo final -208 Osmo initial ), which is a rigorous measure of whole-body hydration in vertebrates (Peterson, 209 2002). For these assessments, females were bled at the onset and at the end of the water 210 restriction period using a standard protocol [START_REF] Meylan | The effect of transdermal corticosterone 609 application on plasma corticosterone levels in pregnant Lacerta vivipara[END_REF]. Blood samples 211 (ca. 40 µL whole blood) were collected from the post-orbital sinus and centrifuged for 5 min 212 at 11,000 rpm. Plasma was then separated from the blood cells and kept frozen at -30°C in 213 airtight tubes until analyses were performed. Plasma osmolality (± 1 mOsm.kg -1 ) was 214 determined using a vapour pressure osmometer (model 5500, Wescor, Logan, UT, USA) 215 according to the protocol previously established in this species (Dupoué et al., 2017a). Before 216 analyses, plasma was diluted (1:1) in a physiological serum (304 mOsm.kg -1 ) so that plasma predicted by their body size [START_REF] Bonnet | Measures of reproductive allometry are 493 sensitive to sampling bias[END_REF]. We used SVL as denominator in this linear 227 regression instead of post-laying BM because i) SVL is fixed and less biased than post-laying 228 BM [START_REF] Bonnet | Measures of reproductive allometry are 493 sensitive to sampling bias[END_REF], ii) SVL-adjustments might better explain reproductive changes 229 than BM-adjustments (Dupoué & Lourdais, 2014;Foucart et al., 2014), and iii) because SVL 230 was a better predictor of female clutch mass (r 2 = 0.34) than post-laying BM (r 2 = 0.21). We 231 calculated the incubation time as the day difference between hatching and oviposition dates.

232

We determined hatching success as the number of live juveniles within the clutch against 233 stillborn and undeveloped eggs. All alive offspring were then counted, weighed (BM, ± 1 234 mg), measured (SVL, ± 1 mm), and sexed by counting ventral scales on the medioventral We examined whether female dehydration rate could be further related to treatment 267 timing and environmental conditions. For each response variable (ΔBM or ΔOsmo), we used 268 the Akaike information criterion corrected for small sample size [AICc, package 269 AICcmodavg [START_REF] Mazerolle | AICcmodavg: Model selection and multimodel inference based on 607 (Q)AIC(c)[END_REF]], to compare a set of models with different environmental 270 measures (Table S3: all females; Table S4: gravid females only). In all models, population 271 was set as a random factor to account for the non-independence of females within population. S3), and iv) interactive models with initial value, 277 SVL, reproductive status, hydric treatment, first and second order interactive effects between 278 hydric treatment, reproductive status and each environmental condition treated separately 279 (altitude, water access, T min and T max , see Table S3). In gravid females specifically, we used a 280 similar procedure to test the effects of initial physiological state, SVL, treatment timing, 281 hydric treatment, environmental conditions and their interaction on ΔBM and ΔOsmo (see all 282 models in Table S4). In some models, altitude was treated as explanatory covariates in 283 addition with SVL or treatment timing, a procedure that may induce multicollinearity given 284 the positive correlation between those (respectively: t 6.1 = 3.1, p = 0.022, t 106.0 = 6.9, p < 285 0.001). Despite this relationship, the Pearson's correlation coefficients between SVL and 286 altitude (r = 0.32) or treatment timing and altitude (r = 0.55) were below the threshold when 287 multicollinearity bias may occur (r = 0.70) [START_REF] Dormann | Collinearity: A review of methods to deal with 505 it and a simulation study evaluating their performance[END_REF].

288

We used similar approaches to determine whether reproductive performance was 289 correlated with treatment timing and whether it differed between hydric treatments, alone, in 290 addition to, or in interaction with environmental conditions (altitude, water access, and T min S5). In all models, population was set as a 292 random factor. Oviposition date (Julian date), RCM and incubation time were analysed with 293 linear mixed models, and hatching success was analysed with mixed-effect logistic 294 regressions including a logit link and binomial error term (number of viable versus number of 295 failed offspring). Offspring SVL and BM were analysed with similar linear mixed models, 296 with offspring sex as an additive or interactive factor (see all models in Table S5) and random 297 factors included both the mother identity and population to account for non-independence 298 between siblings and within populations.

299

Results

300

Physiological responses to water restriction 301

At the onset of experiment, BM ini was higher in gravid than non-gravid females (t 135.6 = -2.6, 302 p = 0.010), irrespective of treatment affiliation (control: 3.200 ± 0.087, water restriction 3.192 303 ± 0.085 g, t 132.9 = -0.5, p = 0.632). Osmo ini did not differ neither between reproductive statuses 304 (t 135.0 = -0.5, p = 0.635) nor between treatment affiliation (control: 319.1 ± 2.2, water 305 restriction 319.4 ± 2.7 mOsm.kg -1 , t 129.4 = 0.2, p = 0.845).

306

At the end of exposure to hydric treatment, changes in body mass (ΔBM) were 307 positively correlated with BM ini (t 134.0 = 4.3, p < 0.001) and lower in water restricted females 308 compared to controls (t 134.0 = -5.5, p < 0.001, Fig. 2A), irrespective of their SVL and 309 reproductive state (Table S3). In gravid females, ΔBM was positively correlated with RCM 310 (t 105.0 = 3.1, p = 0.003, Fig. 2B), and negatively correlated to treatment timing (t 105.0 = -2.1, p 311 = 0.041, Fig. 2C), irrespective of hydric treatment (Table S4). At the same time, the changes S3). In

318

Females from lower altitude populations exhibited higher sensitivity to water 319 restriction since they experienced a greater loss of BM (t 105.0 = 2.9, p = 0.005, Fig. 3A) and 320 higher increases in plasma osmolality (t 105.0 = -3.2, p = 0.002, Fig. 3B) than those from 321 highlands, while no significant altitudinal variation of hydration state occurred in control 322 females (both,p > 0.800,Fig 3A & 3B), irrespective of reproductive status (Table S3).

323

Effects of water restriction on reproductive output

324

Water restriction had no effect on reproductive performance, including oviposition date (mean 325 ± SE, control mothers: 29 th June ± 1 day, water-restricted mothers: 30 th June ± 1 day, t 100.3 = 326 0.8, p = 0.415), clutch mass (control mothers: 1.366 ± 0.059 g, water-restricted mothers: 327 1.370 ± 0.065 g, t 102.7 = 0.0, p = 0.967), RCM (control mothers: 0.005 ± 0.044, water-328 restricted mothers: -0.005 ± 0.055, t 102.6 = -0.1, p = 0.891), incubation time (control mothers: 329 19.0 ± 0.2 days, water-restricted mothers: 19.1 ± 0.2 days, t 73.0 = 0.7, p = 0.479), hatching 330 success (control mothers: 52.3 ± 5.4%, water-restricted mothers: 59.3 ± 5.8%, z = 1.7, p = 331 0.097), nor on offspring SVL (control mothers: 20.07 ± 0.11 mm, water-restricted mothers: 332 20.19 ± 0.10 mm, t 72.5 = 0.5, p = 0.632) or BM (control mothers: 225.1 ± 2.7 mg, water-333 restricted mothers: 229.2 ± 2.7 mg, t 69.6 = 0.5, p = 0.615). Female SVL did not influence 334 oviposition date, RCM, hatching success, and offspring SVL and BM (Table S5) but it was 335 negatively correlated with incubation time (t 78.0 = -2.1, p = 0.037).

336

Effects of environmental conditions on reproductive output

337

Regarding the effects of environmental conditions, we found that altitude and treatment 338 timing significantly explained variation in reproductive performance (Table S5). Gravid females from lower altitude populations laid eggs earlier than those from higher altitude (t 106.0 340 = 8.2, p < 0.001, Fig. 4A), while the incubation time decreased with altitude (t 78.0 = -4.3, p < 341 0.001, Fig. 4B). RCM was negatively correlated with treatment timing (t 106.0 = -5.7, p < 342 0.001), so that females laying eggs sooner also had greater reproductive effort than those with 343 late oviposition (Fig. 4C). Hatching success was negatively correlated with treatment timing 344 (z = -3.0, p = 0.002, Fig. 4D) and interactively impacted by mother treatment and altitude (z = 345 -4.1, p < 0.001), since it decreased along altitude in clutches from water-restricted mothers (z 346 = -3.2, p = 0.002, Fig. 4E), while slightly increasing in those from control mothers (z = 2.1, p 347 = 0.033, Fig. 4E). Both morphometric measures of offspring (SVL and BM) were positively 348 correlated with altitude (offspring SVL: t 7.4 = 4.0, p = 0.005; offspring BM: t 7.1 = 3.6, p = 349 0.008), irrespective of mother treatment (Table S5). In addition, female offspring had longer 350 SVL than males (t 289.3 = 6.0, p < 0.001, Fig 4F ), and sex differences in BM depended upon 351 mother treatment (interaction term: t 269.5 = -3.1, p = 0.002). That is, daughters of control 352 mothers had lower BM than their brothers (t 268.6 = -2.9, p = 0.004, Fig. 4G), while there was 353 no difference between offspring of water-restricted mothers (t 269.3 = 1.5, p = 0.137, Fig. 4G).

354

Discussion

355

Following a two-week water restriction period, both gravid and non-reproductive females 356 responded the same way, suggesting no additional hydric cost of reproduction [sensu 357 [START_REF] Lourdais | Hydric 'costs' of reproduction: pregnancy 597 increases evaporative water loss in the snake Vipera aspis[END_REF]]. This was further confirmed by an absence of relationship between 358 dehydration rate and both the relative reproductive effort and treatment timing. These results

359

likely highlight low water requirements from embryos at early developmental stages, and 360 contrast with recent findings in an oviparous snake, where water restriction applied during the 361 entire gravidity period (three weeks) did affect females' hydration state and resulted in lower 362 egg mass [START_REF] Brusch | Muscles provide an internal water 495 reserve for reproduction[END_REF]. Interestingly, we found that independently of reproductive state, female physiological 364 responses to water restriction were shaped by altitude, while we expected habitat water access 365 or temperature measures of a population to better reflect phenotypic variation in water balance 366 regulation [START_REF] Guillon | Microclimate preferences correlate with 554 contrasted evaporative water loss in parapatric vipers at their contact zone[END_REF][START_REF] Cox | Evolutionary shifts in habitat aridity predict evaporative water 498 loss across squamate reptiles[END_REF]Dupoué et al., 2017a) and may therefore represent a better descriptor of annual conditions than punctual measures of 375 temperatures. Here, females from lower altitudes faced higher dehydration rates probably 376 because they were exposed to water restriction when embryos were more developed with 377 higher water needs [START_REF] Lourdais | Embryonic water uptake during pregnancy is 600 stage-and fecundity-dependent in the snake Vipera aspis[END_REF]. In addition, females at high altitudes may be 378 locally adapted and less permeable to water loss due to body shape [e.g., lower surface-to-379 volume ratio, (Dupoué et al., 2015b)], or more prompt to initiate water-saving strategies (e.g., 380 lower thermal preferences), to limit evaporative water loss and dehydration risk (Köhler et al., 381 2011;Rozen-Rechels et al. in press). In support of this last hypothesis, we recently 382 documented thermal preference negatively correlated with altitude in females from the exact 383 same populations (Trochet et al., 2018).

384

Altitude also correlated with oviposition date (sooner at lower altitudes) and 385 incubation duration (faster at higher altitudes), consistently with expectations and previous 386 evidence on phenology variation along elevation gradients [START_REF] Heulin | Demography of a bimodal reproductive 568 species of lizard (Lacerta vivipara): survival and density characteristics of oviparous 569 populations[END_REF]Rodríguez-387 Díaz & Braña, 2012;[START_REF] Rutschmann | Climate and habitat interact to shape 638 the thermal reaction norms of breeding phenology across lizard populations[END_REF] [START_REF] Packard | The physiological and ecological importance of water to embryos of 612 oviparous reptiles[END_REF] 395 so that this result was surprising. Due to the orographic effects of altitude, females from low 396 altitudes regularly face periods of more severe water restriction (summer droughts) compared 397 to those from high altitudes with more frequent rain episodes. This contrast might have 398 directly resulted in some degree of local adaptation with enhanced reproductive fitness for 399 low altitude females in the drier conditions (water restriction treatment) and in high altitude 400 females in the wetter conditions (control treatment). Additionally, greater hatching success in 401 water-restricted females from low altitudes might have indirectly resulted from locally 402 adapted thermoregulation. As stated before, females from low altitude populations have 403 higher preferred body temperatures (Trochet et al., 2018). In this species, small differences in 404 thermoregulation during gravidity may strongly impact hatching success (Foucart et al.,405 2018). Surviving offspring from higher altitudes had larger body size and body mass than 406 those from lower altitudes. Overall, daughters were longer than sons as classically 407 documented in this species [START_REF] Lecomte | Sex identification in juveniles of Lacerta vivipara[END_REF]Le Galliard et al., 2006;Dupoué et al., 408 2019), and sex differences in mass depended upon mother treatment. That is, daughters from 409 control mothers had lower body mass than sons, while offspring from water-restricted 410 mothers had similar mass. Altogether, our results therefore suggest multi-level trade-offs 411 since females from low altitudes laid eggs sooner and despite prolonged egg incubation time 412 in standard conditions, they had greater hatching success but produced smaller offspring. These trade-offs were relatively independent from punctual dehydration but they were rather 414 shaped by long-term acclimation or local adaptation to altitude.

415

In the Western viviparous lineages of the common lizard, population extinction risk is 416 increasing at low altitudes where lizards experience higher temperatures (Massot, Clobert, & 417 Ferrière, 2008;Sinervo et al., 2010) and associated costs (Dupoué et al., 2017b(Dupoué et al., , 2018b)).

418

Viviparous species are generally predicted to be more vulnerable to climate change than 419 oviparous ones (Sinervo et al., 2010(Sinervo et al., , 2018)). In support of this hypothesis, the oviparous 420 common lizard remains present (in low abundance) at sea level and only in humid landscapes 421 (peat bogs, forest marshes) that support its presence [START_REF] Berroneau | Atlas des amphibiens et reptiles d'Aquitaine (C Nature[END_REF]. Although causal 422 factors remain elusive, we showed here that water restriction early in gravidity had relatively 423 low physiological impacts on the oviparous form and this contrasts with previous findings in 424 the viviparous form (Dupoué et al., 2018a). In fact, our results suggest that the oviparous 425 form at low altitudes might even gain in fitness (hatching success) when exposed to water 426 restriction. Future work is now critically needed to experimentally manipulate both 427 temperature and water to measure the adaptive significance and long-term response in this 428 bimodal species.

429

To conclude, the evolution of reproductive strategies constitutes the core part of life-430 history theory (Stearns, 1992). Given their outstanding variability and flexibility in 431 reproductive modes, squamates remain key models to investigate the causes and 432 consequences of transitioning to viviparity [START_REF] Blackburn | Squamate reptiles as model organisms for the evolution of viviparity[END_REF]Laird, Thompson, & 433 Whittington, 2019). Here, we repeated an experimental procedure in both forms of a 434 reproductively bimodal species. Although some fitness consequences may also appear over form (35 to 3 days before parturition). We hypothesized water constraints to be relatively low 678 in oviparous females given that most of embryo water demand occurs during somatic growth 679 [START_REF] Lourdais | Embryonic water uptake during pregnancy is 600 stage-and fecundity-dependent in the snake Vipera aspis[END_REF], once oviparous females have laid their eggs. q q q q *** ( ) q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q RCM (residuals) q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q -1.6 -0.8 0 0.8 -500 0 500 1000 B) q q Control Water-restricted q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q Timing (days before oviposition) q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q 0 10 20 30 q q q q *** ( ) q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q RCM (residuals) q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q E) -1.6 -0.8 0 0.8 -50 0 50 100 q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q Timing (days before oviposition) q q q q q q q q q q q q q qq q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q F) 0 q q q q q q q q q q q q Altitude (m) q q q q q q q q q q q q Page 33 of 57 10-Jul q q q q q q q q q qq q q q Control Water-restricted Altitude (m) Incubation time (days) B) 1000 1200 1400 1600 17 18 19 20 q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q Timing (days before oviposition) RCM (residuals) q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q C) 0 10 20 30 -1.4 -0.7 0 0.7 q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q qq q q q q q q q q q q q q q q q q q q q q q q Timing (days before oviposition) Hatching success (%) q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q D) 0 10 20 30 0 25 50 75 100 q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q q Altitude (m) Hatching success (%) q q q q q q q q q q q q Offspring sex SVL (mm)

Biological

F M 19.4 20.2 21 F) q q q q n.s. { } Offspring sex BM (mg) F M 218 230 242 G) q q q q ** { } F o r P e e r
F M 23 M M M M M M - - - - F 143 4 24 M M M M M M - - - - M 4 75 25 M M M M M M - - - - 26 M M M M M M - - - - 27 M M M M M M F F F F 28 M M M M M F F F F F 29 - - - - F F F F F F 30 - - - - F F F F F F 31 - - - - F F F F F F 32 - - - - F F F F F F

  -offs, by shaping the allocation of resources within and between generations,

  clutches (range = 1 to 9 eggs) were weighed (± 1 mg) and placed in 194 individual plastic cups on water-saturated vermiculite to maintain hydric conditions and 195 incubated at T set = 25°C to optimise incubation time without risk of overheating for embryo 196 development

197(

  Novital Covatutto Eco, Italy) and we randomly distributed the clutches in the three 198 incubators (n = 35 per incubator) that remained within a 1°C range of observed temperatures 199 (mean ± SD, incubator 1: T obs = 24.38 ± 0.26°C; incubator 2: T obs = 24.05 ± 0.39°C; incubator 200 3: T obs = 24.63 ± 0.30°C). Juveniles were released within 3 days post-hatching in their 201 respective populations. 202 Female water balance 203

  effects of water restriction on different reproductive traits. We checked 221 females daily to compare oviposition date. For each female we subtracted the oviposition date 222 from the last day in hydric treatment to calculate the treatment timing. As an index of 223 reproductive effort, we estimated size-adjusted relative clutch mass (RCM) at oviposition 224 (i.e., residuals from the linear relationship between clutch mass and body size, CM -SVL: t 103 225 = 7.2, p < 0.001, Fig. S1) to determine how much females deviate from the reproductive effort 226

272

  Our set of models included i) a null model with only the intercept and random factor, ii) 273 simple models (initial values of BM, SVL, osmolality, reproductive status or hydric treatment 274 alone), iii) additive models with initial value, SVL, reproductive status, hydric treatment and 275 additive effects of each environmental conditions (altitude, water access, and T min and T max 276 treated separately; see all models in Table

  treated separately; see all models in Table

312

  in plasma osmolality (ΔOsmo) were negatively correlated with Osmo ini (t 133.2 = -8.3, p < 313 0.001) and significantly increased following water restriction compared to control conditions 314 (t 129.2 = 6.6, p < 0.001, Fig 2D) irrespective of their SVL and reproductive state (Table

  ΔOsmo was neither correlated with RCM (t 104.9 = -0.8, p = 0.410, Fig.2E), 316 nor with treatment timing (t 105.0 = -0.1, p = 0.957, Fig 2F), in both hydric treatments (

  435time, the lack of changes in females physiology depending on reproductive effort suggest that 436 they did not pay immediate water costs contrary to their viviparous relative(Dupoué et al., prolonged egg retention when evolving in water limiting environments.439Additionally, water constraints may also favour a reverse transition from viviparity to 440 oviparity, if the hydric costs of reproduction become too high[START_REF] Lourdais | Hydric 'costs' of reproduction: pregnancy 597 increases evaporative water loss in the snake Vipera aspis[END_REF] Dupoué 441 et al., in press). In support of this innovative hypothesis, phylogenomic analyses unravelled 442 that the Western lineages populations sampled here likely originated from a viviparous 443 ancestor and evolved back to an oviparous reproductive mode[START_REF] Recknagel | Common lizards break Dollo's law of 626 irreversibility: Genome-wide phylogenomics support a single origin of viviparity and re-627 evolution of oviparity[END_REF] Horreo 444 et al., 2020). Hence, egg-laying might have evolved during the last Pleistocene and Holocene 445 as an adaptation to high habitat aridity. Our study therefore calls future work to test this 446 hypothesis and include water constraints when studying the environmental drivers of 447 reproductive strategies. 448 'Office Nationale des Forêts', the 'Parc National des Pyrénées' and the region Occitanie for 452 allowing us to sample lizards. We thank Pr Daniel G. Blackburn and three anonymous 453 reviewers for greatly improving the early drafts of this manuscript. the financial and technical support from the Centre National de la 458 Recherche Scientifique (CNRS), and was funded by the Interreg Poctefa under the project 459 'Ectopyr' (EFA031/15 to FA), by the Agence Nationale de la Recherche under the Aquastress' project (2018-1R20214 to OL). AD was financially supported by the 462 Australian Research Council (DP180100058 to A/Prof Anne Peters) at the time of writing. P, Rozen-Rechels D, et al. Water availability and temperature induce 510 changes in oxidative status during pregnancy in a viviparous lizard. Functional Ecology. in 511 press 512 Dupoué A, Brischoux F, Angelier F, et al. 2015a. Intergenerational trade-off for water may 513 induce a mother-offspring conflict in favour of embryos in a viviparous snake. Functional 514 Ecology 29: 414-422. 515 Dupoué A, Le Galliard JF, Josserand R, et al. 2018a. Water restriction causes an 516 intergenerational trade-off and delayed mother-offspring conflict in a viviparous lizard. 517 Functional Ecology 32: 676-686. 518 Dupoué A & Lourdais O. 2014. Relative reproductive effort drives metabolic changes and 519 maternal emaciation during pregnancy in a viviparous snake. Journal of Zoology 293: 49-56. 520 Dupoué A, Lourdais O, Meylan S, et al. 2019. Some like it dry: Water restriction overrides 521 heterogametic sex determination in two reptiles. Ecology and Evolution 9: 6524-6533. 522 Dupoué A, Rutschmann A, Le Galliard JF, et al. 2017a. Water availability and 523 environmental temperature correlate with geographic variation in water balance in common 524 lizards. Oecologia 185: 561-571. 525 Dupoué A, Rutschmann A, Le Galliard JF, et al. 2017b. Shorter telomeres precede 526 population extinction in wild lizards. Scientific Reports 7: 16976. 527 Dupoué A, Rutschmann A, Le Galliard JF, et al. 2018b. Reduction of baseline 528 corticosterone secretion correlates with climate warming and drying across wild lizard 529 populations. Journal of Animal Ecology 87: 1331-1341.

  -de-la-Cruz F, Miles DB, et al. 2010. Erosion of lizard diversity by 649 climate change and altered thermal niches. Science 328: 894-899. 650 Sinervo B, Miles DB, Wu Y, et al. 2018. Climate change, thermal niches, extinction risk and 651 maternal-effect rescue of toad-headed lizards, Phrynocephalus, in thermal extremes of the 652 Arabian Peninsula to the Qinghai-Tibetan Plateau. Integrative Zoology 13: 450-470. 653 Stearns SC. 1992. The evolution of life histories. Oxford: Oxford University Press. 654 Stewart JR & Blackburn DG. 2015. Viviparity and placentation in lizards. In: Sever D, 655 Aldridge R, eds. Reproductive Biology and Phylogeny of Snakes. Enfield, New Hampshire, 656 USA, 119-181. 657 Surget-Groba Y, Heulin B, Guillaume CP, et al. 2006. Multiple origins of viviparity, or 658 reversal from viviparity to oviparity? The European common lizard (Zootoca vivipara, 659 Lacertidae) and the evolution of parity. Biological Journal of the Linnean Society 87: 1-11. 660 Thorpe RS & Baez M. 1993. Geographic variation in scalation of the lizard Gallotia stehlini 661 within the island of Gran Canaria. Biological Journal of the Linnean Society 48: 75-87. 662 Trivers RL. 1974. Parent-offspring conflict. American Zoologist 14: 249-264. 663 Trochet A, Dupoué A, Souchet J, et al. 2018. Variation of preferred body temperatures 664 along an altitudinal gradient: A multi-species study. Journal of Thermal Biology 77: 38-44. 665 Zeh DW & Zeh JA. 2000. Reproductive mode and speciation: the viviparity-driven conflict 666 hypothesis. BioEssays 22: 938-946.

Figure 1 .

 1 Figure legends

  677

Figure 2 .<Figure 3 .Figure 4 .

 234 Figure 2. Examination of physiological responses to water restriction in gravid and non-

  Journal of the Linnean Society Biological Journal of the Linnean Society F o r P e e r R e v i e w

Figure S2 .

 S2 Figure S2. Positive relationship between female body size (snout-vent length -SVL) and altitude (t 6.4 = 3.1, p = 0.022). Females had similar SVL between the affiliation to waterrestriction (filled circles) or control conditions (open circles) (t 129.4 = -0.1, p = 0.908), irrespective of the altitude of the population of origin (t 129.3 = 0.1, p = 0.929).

Table 317 S4 ) .

 317)

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,

	Mother-offspring Fabien Aubret Mother-offspring
	F
	o r
	P
	e e r
	R e v R e v
	i e w i e w

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert, Fabien Aubret

Table S1 .

 S1 Localizations, altitude, climatic conditions and water access (permanent in peatbog type habitat, periodic in dry meadow) in the 6 natural populations of the oviparous form of common lizard (Z. vivipara) from the Pyrenees Mountain range. Table shows the mean ± SE of environmental temperatures (minimal: T min and maximal: T max ) recorded between June 30 th and July 25 th , 2018. See text for details.

	Page 37 of 57	Biological Journal of the Linnean Society
	Mother-offspring
	Population Col des Marrous 42°57'44.26"N ; 1°26'40.36"E 990 GPS localization Altitude Water access T min Periodic 14.75 ± 0.20 17.50 ± 0.22 T max Lac de Bethmale 42°51'43.62"N ; 1°05'03.77"E 1054 Permanent 13.06 ± 0.19 16.77 ± 0.31 Col de la Core 42°51'33.37"N ; 1°06'20.12"E 1405 Periodic 11.56 ± 0.30 19.82 ± 0.65 Etang de Lers 42°48'32.06"N ; 1°22'32.56"E 1246 Permanent 13.75 ± 0.20 17.21 ± 0.30 Aulus 42°46'57.47"N ; 1°23'11.68"E 1400 Permanent 14.50 ± 0.21 19.81 ± 0.39 Col d'Agnes 42°47'39.69"N ; 1°22'27.63"E 1580 Periodic 12.33 ± 0.19 17.91 ± 0.47 F o r P e e r R e v i e w
		Biological Journal of the Linnean Society

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert, Fabien Aubret

Table S2 .

 S2 Sex determination table using scale counting along medioventral (left and right side) as previously described[START_REF] Lecomte | Sex identification in juveniles of Lacerta vivipara[END_REF].

	Left table reports the number of ventral scales in reproductive adults (147 females and 79 males) with obvious secondary sexual traits
	phenotypes. Right table illustrates that scale counting matched phenotypic sex with high determination success (96.5%).
		Sex determination using number of ventral scales	N individuals with sex determined
	left	right	23 24 25 26 27 28 29 30 31 32	scales	phenotypic

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Biological Journal of the Linnean Society	Page 38 of 57
	Mother-offspring Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
	Fabien Aubret
	F
	o r
	P
	e e r
	R e v
	i e w
	Biological Journal of the Linnean Society

Table S3 .

 S3 AICc based model selection in all females (gravid and non-reproductive) comparing a null model (intercept only) to models testing relationships between female dehydration indexes (changes in body mass ΔBM and changes in plasma osmolality ΔOsmo)

	Mother-offspring
	and reproductive status, hydric treatment and environmental conditions. Models included the
	initial physiological value (BM ini or Osmo ini ), SVL, altitude and temperature indexes (T min and
	T max ) treated as linear co-variates, while reproductive status (gravid versus non-reproductive)
	hydric treatment (control versus water restriction) and water access (permanent versus
	periodic) were treated as fixed factors. Population was set as a random factor in all models to
	account for intra-population variation and non-independence. Highlighted models (bolded
	characters) are those explaining the greatest variation in response variables. F
	Physiological responses ΔBM	Models BM ini + status + treatment + status * o r treatment P BM ini + status + treatment + altitude + treatment * altitude BM ini + status + treatment BM ini + treatment + status + altitude + e e r	k 7 8 6	AICc 8.64 9.28 9.64	ΔAICc 0.00 0.64 1.00	w i 0.13 0.10 0.08	LogLike 3.13 3.94 1.51
		treatment * status + treatment * altitude + status * altitude BM ini + status + treatment + water access + treatment * water access BM ini + treatment + status + altitude + R 10 8 8 e v 10.20 9.73 10.51 treatment * status BM ini + treatment + status + Tmax + treatment * status 8 10.53 BM ini + treatment + status + water access + treatment * status 8 10.77 i e w	1.09 1.57 1.87 1.90 2.14	0.08 0.06 0.05 0.05 0.05	6.03 3.48 3.32 3.31 3.19
		BM ini + treatment	5	10.80	2.17	0.05	-0.17
		BM ini + treatment + status + Tmin + treatment * status	8	10.85	2.22	0.04	3.15
		SVL + BM ini + status + treatment + status * treatment	8	10.89	2.26	0.03	3.13
		BM ini + treatment + status + altitude +
		treatment * status + treatment * altitude + status * altitude + treatment * status *	11	11.03	2.39	0.04	6.57
		altitude
		BM ini + status + treatment + Tmax	7	11.31	2.67	0.04	1.79
		BM ini + status + treatment + altitude	7	11.37	2.73	0.03	1.76
		BM ini + status + treatment + Tmin + treatment * Tmin	8	11.53	2.89	0.03	2.81
		SVL + BM ini + status + treatment + altitude + treatment * altitude	9	11.56	2.92	0.02	3.95

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
	Fabien Aubret
	BM ini + status + treatment + water access	7	11.78	3.14	0.03	1.55
	SVL + BM ini + status + treatment	7	11.86	3.22	0.02	1.52
	BM ini + status + treatment + Tmin	7	11.86	3.23	0.03	1.51
	SVL + BM ini + treatment + status + altitude
	+ treatment * status + treatment * altitude	11	12.06	3.43	0.02	6.05
	+ status * altitude
	SVL + BM ini + status + treatment + water access + treatment * water access	9	12.48	3.84	0.01	3.49
	BM ini + treatment + status + water access +
	treatment * status + treatment * water	10	12.61	3.98	0.02	4.59
	access + status * water access
	SVL + BM ini + treatment + status + altitude + treatment * status	9	12.80	4.16	0.01	3.33
	SVL + BM ini + treatment + status + Tmax + F treatment * status SVL + BM ini + treatment BM ini + treatment + status + Tmin + o r	9 6	12.83 12.99	4.19 4.36	0.01 0.01	3.31 -0.17
	treatment * status + treatment * Tmin + status * Tmin P	10	13.00	4.36	0.02	4.40
	BM ini + treatment + status + Tmax + status * Tmax SVL + BM ini + treatment + status + water access + treatment * status e e r SVL + treatment + status + Tmin + treatment * status SVL + BM ini + treatment + status + altitude + treatment * status + treatment * altitude + status * altitude + treatment * status * altitude R 12 8 9 9 e v 13.01 13.07 13.14 13.44	4.37 4.43 4.50 4.81	0.02 0.01 0.01 0.01	2.07 3.19 3.16 6.57
	SVL + BM ini + status + treatment + Tmax BM ini + status + treatment + Tmax + treatment * Tmax SVL + BM ini + status + treatment + altitude 8 8 8 i 13.57 13.57 13.62 e w	4.94 4.94 4.99	0.01 0.01 0.01	1.79 1.79 1.76
	BM ini + treatment + status + altitude + status * altitude	8	13.63	4.99	0.01	1.76
	SVL + BM ini + status + treatment + Tmin + treatment * Tmin	9	13.78	5.15	0.01	2.83
	BM ini + treatment + status + Tmin + status * Tmin	8	13.80	5.17	0.01	1.67
	BM ini + treatment + status + water access + status * water access	8	13.92	5.28	0.01	1.62
	SVL + BM ini + status + treatment + water access	8	14.04	5.40	0.01	1.56
	SVL + BM ini + status + treatment + Tmin	8	14.12	5.48	0.01	1.52
	BM ini + treatment + status + water access +
	treatment * status + treatment * water access + status * water access + treatment	11	14.82	6.19	0.01	4.67
	* status * water access

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
		Fabien Aubret
		BM ini + treatment + status + Tmax +
		treatment * status + treatment * Tmax +	10	14.89	6.25	0.01	3.45
		status * Tmax
		SVL + BM ini + treatment + status + water
		access + treatment * status + treatment *	11	14.97	6.33	0.00	4.60
		water access + status * water access
		SVL + BM ini + treatment + status + Tmax + status * Tmax	9	15.29	6.66	0.00	2.08
		SVL + BM ini + treatment + status + Tmin +
		treatment * status + treatment * Tmin +	11	15.33	6.69	0.00	4.42
		status * Tmin
		BM ini + treatment + status + Tmin +
		treatment * status + treatment * Tmin +	11	15.35	6.72	0.00	4.41
		status * Tmin + treatment * status * Tmin
		SVL + BM ini + status + treatment + Tmax + treatment * Tmax	9	15.87	7.23	0.00	1.79
		SVL + BM ini + treatment + status + altitude + status * altitude F	9	15.92	7.28	0.00	1.77
		SVL + BM ini + treatment + status + Tmin + status * Tmin SVL + BM ini + treatment + status + water o r access + status * water access BM ini + treatment + status + Tmax + treatment * status + treatment * Tmax + status * Tmax + treatment * status * Tmax P SVL + BM ini + treatment + status + water access + treatment * status + treatment * water access + status * water access + e e r	9 9 11 12	16.10 16.20 16.34 17.22	7.47 7.57 7.71 8.58	0.00 0.00 0.00 0.00	1.67 1.62 3.91 4.68
		treatment * status * water access SVL + BM ini + treatment + status + Tmax + treatment * status + treatment * Tmax + status * Tmax SVL + BM ini + treatment + status + Tmin + treatment * status + treatment * Tmin + status * Tmin + treatment * status * Tmin R 11 12 e v 17.26 17.71 SVL + BM ini + treatment + status + Tmax + treatment * status + treatment * Tmax + status * Tmax + treatment * status * Tmax 12 18.76 i e w	8.62 9.08 10.12	0.00 0.00 0.00	3.45 4.43 3.91
		BM ini + status	5	34.61	25.97	0.00	-12.07
		BM ini	4	34.77	26.13	0.00	-13.23
		SVL + BM ini + status	6	36.80	28.16	0.00	-12.07
		SVL + BM ini	5	36.92	28.28	0.00	-13.22
		SVL	4	40.93	32.29	0.00	-16.31
		Null	3	51.26	42.63	0.00	-22.54
	ΔOsmo	Osmo ini + status + treatment + altitude + treatment * altitude	8	1239.24	0.00	0.45	-611.04
		SVL + Osmo ini + status + treatment + altitude + treatment * altitude	9	1241.52	2.29	0.11	-611.04
		Osmo ini + status + treatment + altitude	7	1242.15	2.91	0.11	-613.63

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
	Fabien Aubret
	Osmo ini + treatment + status + altitude +
	treatment * status + treatment * altitude +	10 1243.16	3.92	0.06	-610.69
	status * altitude
	Osmo ini + treatment + status + altitude + status * altitude	8	1243.73	4.49	0.05	-613.29
	Osmo ini + treatment	5	1243.76	4.52	0.05	-616.65
	Osmo ini + status + treatment + Tmax + treatment * Tmax	8	1243.82	4.59	0.05	-613.34
	Osmo ini + status + treatment + Tmax	7	1244.27	5.03	0.04	-614.69
	SVL + Osmo ini + status + treatment + altitude	8	1244.38	5.15	0.03	-613.62
	Osmo ini + treatment + status + altitude + treatment * status	8	1244.40	5.17	0.03	-613.63
	Osmo ini + status + treatment F Osmo ini + treatment + status + altitude + treatment * status + treatment * altitude + status * altitude + treatment * status * altitude o r	6 11 1245.43 1244.56	5.33 6.19	0.03 0.02	-615.95 -610.63
	SVL + Osmo ini + treatment + status + altitude + treatment * status + treatment * P	11 1245.47	6.24	0.01	-610.66
	altitude + status * altitude SVL + Osmo ini + status + treatment + Tmax + treatment * Tmax SVL + Osmo ini + treatment e e r	9 6	1245.51 1245.70	6.27 6.46	0.01 0.01	-613.03 -616.52
	SVL + Osmo ini + status + treatment + Tmax SVL + Osmo ini + treatment + status + altitude + treatment * status Osmo ini + status + treatment + water access + treatment * water access R e 8 9 8 v 1245.92 1245.95 1246.11 SVL + Osmo ini + status + treatment 7 1246.33 Osmo ini + treatment + status + Tmax + treatment * status 8 1246.50 i e w	6.68 6.71 6.87 7.10 7.26	0.01 0.01 0.01 0.01 0.01	-614.38 -613.25 -614.48 -615.72 -614.67
	Osmo ini + treatment + status + Tmax + status * Tmax	8	1246.50	7.27	0.01	-614.68
	SVL + Osmo ini + treatment + status + altitude + treatment * status	9	1246.68	7.44	0.01	-613.61
	Osmo ini + status + treatment + Tmin	7	1246.71	7.47	0.01	-615.91
	Osmo ini + status + treatment + water access 7	1246.77	7.53	0.01	-615.94
	Osmo ini + status + treatment + status * treatment	7	1246.79	7.55	0.01	-615.95
	Osmo ini + treatment + status + water access + status * water access	8	1246.94	7.70	0.01	-614.89
	SVL + Osmo ini + treatment + status +
	altitude + treatment * status + treatment * altitude + status * altitude + treatment *	12 1247.81	8.57	0.00	-610.61
	status * altitude
	Osmo ini + treatment + status + Tmin + status * Tmin	8	1247.89	8.65	0.01	-615.37

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
	Fabien Aubret
	Osmo ini + treatment + status + water access
	+ treatment * status + treatment * water access + status * water access + treatment	11 1247.99	8.75	0.01	-611.91
	* status * water access
	SVL + Osmo ini + status + treatment + water access + treatment * water access	9	1248.07	8.84	0.00	-614.31
	SVL + Osmo ini + status + Tmax + status * treatment	9	1248.20	8.96	0.00	-614.37
	SVL + Osmo ini + treatment + status + Tmax + status * Tmax	9	1248.21	8.97	0.00	-614.38
	Osmo ini + treatment + status + Tmax +
	treatment * status + treatment * Tmax +	11 1248.33	9.10	0.00	-612.09
	status * Tmax + treatment * status * Tmax
	Osmo ini + treatment + status + Tmax +
	treatment * status + treatment * Tmax +	10 1248.38	9.15	0.00	-613.30
	status * Tmax
	SVL + Osmo ini + status + treatment + Tmin 8 F	1248.44	9.21	0.00	-615.65
	SVL + Osmo ini + status + treatment + water access o r SVL + Osmo ini + status + treatment + status * treatment Osmo ini + treatment + status + water access P	8 8	1248.56 1248.59	9.33 9.36	0.00 0.00	-615.71 -615.72
	+ treatment * status + treatment * water access + status * water access Osmo ini + treatment + status + Tmin + treatment * status e e r Osmo ini + status + treatment + Tmin + treatment * Tmin SVL + Osmo ini + treatment + status + water access + status * water access Osmo ini + treatment + status + water access + treatment * status R 10 1248.66 8 1248.97 8 1248.97 9 1249.01 8 1249.03 e v SVL + Osmo ini + treatment + status + Tmin + status * Tmin 9 1249.58 SVL + Osmo ini + treatment + status + Tmax + status * Tmax 12 1249.82 SVL + Osmo ini + treatment + status + i e w	9.42 9.73 9.73 9.78 9.79 10.34 10.59	0.00 0.00 0.00 0.00 0.00 0.00 0.00	-613.43 -615.91 -615.91 -614.78 -615.94 -615.06 -611.62
	Tmax + treatment * status + treatment *	11 1250.19	10.95	0.00	-613.01
	Tmax + status * Tmax
	SVL + Osmo ini + treatment + status + water
	access + treatment * status + treatment * water access + status * water access +	12 1250.36	11.12	0.00	-611.89
	treatment * status * water access
	SVL + Osmo ini + treatment + status + Tmin + treatment * status	9	1250.73	11.49	0.00	-615.64
	SVL + Osmo ini + status + treatment + Tmin + treatment * Tmin	9	1250.74	11.51	0.00	-615.65
	SVL + Osmo ini + treatment + status + water access + treatment * status	9	1250.86	11.62	0.00	-615.70
	SVL + Osmo ini + treatment + status + water
	access + treatment * status + treatment *	11 1250.90	11.66	0.00	-613.37
	water access + status * water access
	Osmo ini + treatment + status + Tmin +
	treatment * status + treatment * Tmin +	10 1252.45	13.21	0.00	-615.33
	status * Tmin

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
	Fabien Aubret
	SVL + Osmo ini + treatment + status + Tmin
	+ treatment * status + treatment * Tmin +	11 1254.19	14.95	0.00	-615.01
	status * Tmin
	Osmo ini + treatment + status + Tmin +
	treatment * status + treatment * Tmin +	11 1254.83	15.59	0.00	-615.33
	status * Tmin + treatment * status * Tmin
	SVL + Osmo ini + treatment + status + Tmin
	+ treatment * status + treatment * Tmin +	12 1256.57	17.33	0.00	-615.00
	status * Tmin + treatment * status * Tmin
	Osmo ini	4	1278.83	39.59	0.00	-635.26
	Osmo ini + status	5	1279.89	40.65	0.00	-634.71
	SVL + Osmo ini	5	1280.71	41.47	0.00	-635.12
	SVL + Osmo ini + status Null F SVL o r	6 3 4	1281.50 1319.57 1321.67	42.26 80.33 82.43	0.00 0.00 0.00	-634.42 -656.69 -656.68
	P
	e e r
	R e v
		i e w

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert, Fabien Aubret

Table S4 .

 S4 AICc based model selection in gravid female only comparing a null model (intercept only) to models testing relationships between female dehydration indexes (changes in body mass ΔBM and changes in plasma osmolality ΔOsmo) and environmental conditions. Models included the initial physiological value (BM ini or Osmo ini ), SVL, treatment timing (number of days between laying date and the end of water restriction), altitudes and temperature indexes (T min and T max ) treated as linear co-variates, while hydric treatment (control versus water restriction) and water access (permanent versus periodic) were treated as fixed factors. Population was set as a random factor in all models to account for intrapopulation variation and non-independence. Highlighted models (bolded characters) are those explaining the greatest variation in response variables.

	Mother-offspring
		F
	Physiological responses ΔBM	Models BM ini + timing + treatment + altitude + o r treatment * altitude P BM ini + treatment + altitude + treatment * altitude SVL + BM ini + timing + treatment + altitude + treatment * altitude BM ini + timing + treatment + altitude e e r	k 8 7 9 7	AICc -2.94 -1.09 -0.55 -0.40	ΔAICc 0.00 1.85 2.38 2.54	w i 0.38 0.15 0.09 0.11	LogLike 10.22 8.12 10.22 7.78
		BM ini + timing + treatment BM ini + treatment SVL + BM ini + treatment + altitude + R e 6 5 v treatment * altitude 8 BM ini + timing + treatment + Tmax 7 SVL + BM ini + timing + treatment + altitude 8 i 0.67 0.81 0.82 1.94 1.95 e w	3.61 3.75 3.76 4.88 4.88	0.06 0.06 0.04 0.03 0.02	6.09 4.90 8.34 6.61 7.78
		BM ini + treatment + water access + treatment * water access	7	2.44	5.38	0.03	6.36
		BM ini + treatment + water access	6	2.78	5.71	0.02	5.04
		BM ini + treatment + altitude	6	2.80	5.74	0.02	5.03
		SVL + BM ini + treatment	6	2.80	5.74	0.02	5.03
		BM ini + timing + treatment + water access + treatment * water access	8	2.84	5.78	0.02	7.33
		BM ini + timing + treatment + water access	7	2.93	5.86	0.02	6.11
		BM ini + treatment + Tmax	6	2.95	5.89	0.02	4.95
		BM ini + timing + treatment + Tmin	7	2.96	5.89	0.02	6.10
		BM ini + treatment + Tmin	6	2.96	5.90	0.02	4.95

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
		Fabien Aubret
		SVL + BM ini + timing + treatment	7	2.97	5.91	0.01	6.09
		BM ini + timing + treatment + Tmax + treatment * Tmax	8	3.36	6.29	0.02	7.07
		BM ini + treatment + Tmin + treatment * Tmin	7	3.92	6.86	0.01	5.62
		BM ini + timing + treatment + Tmin + treatment * Tmin	8	4.28	7.22	0.01	6.61
		SVL + BM ini + timing + treatment + Tmax 8	4.29	7.23	0.01	6.61
		SVL + BM ini + treatment + water access + treatment * water access	8	4.56	7.50	0.01	6.47
		SVL + BM ini + treatment + altitude	7	4.71	7.65	0.01	5.22
		SVL + BM ini + treatment + water access	7	4.81	7.75	0.01	5.17
		SVL + BM ini + treatment + Tmax F	7	4.94	7.87	0.01	5.11
		BM ini + treatment + Tmax + treatment * Tmax o r	7	5.00	7.94	0.01	5.08
		SVL + BM ini + treatment + Tmin SVL + BM ini + timing + treatment + water access + treatment * water access P SVL + BM ini + timing + treatment + water access SVL + BM ini + timing + treatment + Tmin e e r	7 9 8 8	5.03 5.23 5.27 5.30	7.96 8.17 8.21 8.24	0.01 0.00 0.00 0.00	5.06 7.33 6.11 6.10
		SVL + BM ini + timing + treatment + Tmax + treatment * Tmax SVL + BM ini + treatment + Tmin + treatment * Tmin SVL + BM ini + timing + treatment + Tmin + treatment * Tmin R e 9 8 9 v SVL + BM ini + treatment + Tmax + treatment * Tmax 8 BM ini + timing 5 27.49 5.75 6.01 6.67 7.02 i e w	8.69 8.94 9.61 9.96 30.42	0.00 0.00 0.00 0.00 0.00	7.07 5.75 6.61 5.24 -8.44
		BM ini	4	28.70	31.64	0.00	-10.15
		SVL + BM ini + timing	6	29.73	32.67	0.00	-8.44
		SVL + BM ini	5	30.59	33.53	0.00	-9.99
		SVL	4	35.52	38.46	0.00	-13.56
		Null	3	42.03	44.96	0.00	-17.89
	ΔOsmo	Osmo ini + treatment + altitude + treatment * altitude	7	961.79	0.00	0.49	-473.32
		SVL + Osmo ini + treatment + altitude + treatment * altitude	8	964.11	2.32	0.12	-473.30
		Osmo ini + timing + treatment + altitude + treatment * altitude	8	964.13	2.34	0.15	-473.32
		Osmo ini + treatment + altitude	6	964.33	2.54	0.14	-475.74

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
	Fabien Aubret
	Osmo ini + timing + treatment + altitude	7	966.50	4.71	0.05	-475.67
	SVL + Osmo ini + timing + treatment + altitude + treatment * altitude	9	966.50	4.71	0.04	-473.30
	SVL + Osmo ini + treatment + altitude	7	966.61	4.83	0.03	-475.73
	Osmo ini + treatment + water access + treatment * water access	7	966.66	4.87	0.04	-475.75
	Osmo ini + treatment + Tmax	6	967.74	5.95	0.02	-477.44
	Osmo ini + treatment	5	967.89	6.10	0.02	-478.64
	Osmo ini + timing + treatment + water access + treatment * water access	8	967.96	6.17	0.02	-475.23
	SVL + Osmo ini + timing + treatment + altitude	8	968.84	7.05	0.01	-475.67
	SVL + Osmo ini + treatment + water access F + treatment * water access Osmo ini + timing + treatment o r Osmo ini + treatment + Tmax + treatment * Tmax SVL + Osmo ini + treatment + Tmax P SVL + Osmo ini + treatment Osmo ini + timing + treatment + Tmax e e r	8 6 7 7 6 7	968.90 969.16 969.51 969.54 969.74 969.83	7.11 7.37 7.72 7.75 7.95 8.04	0.01 0.01 0.01 0.01 0.01 0.01	-475.70 -478.15 -477.18 -477.19 -478.44 -477.34
	Osmo ini + treatment + water access Osmo ini + treatment + Tmin SVL + Osmo ini + timing + treatment + water access + treatment * water access R e 6 6 9 v 969.95 970.07 970.13 SVL + Osmo ini + timing + treatment 7 971.00 Osmo ini + timing + treatment + water access 7 971.38 i e w	8.17 8.28 8.34 9.22 9.60	0.01 0.01 0.01 0.00 0.00	-478.55 -478.61 -475.12 -477.92 -478.11
	Osmo ini + timing + treatment + Tmax + treatment * Tmax	8	971.42	9.63	0.00	-476.96
	SVL + Osmo ini + treatment + Tmax + treatment * Tmax	8	971.43	9.64	0.00	-476.96
	Osmo ini + timing + treatment + Tmin	7	971.45	9.66	0.00	-478.15
	SVL + Osmo ini + timing + treatment + Tmax	8	971.70	9.91	0.00	-477.10
	SVL + Osmo ini + treatment + water access	7	971.88	10.10	0.00	-478.36
	SVL + Osmo ini + treatment + Tmin	7	971.99	10.20	0.00	-478.42
	Osmo ini + treatment + Tmin + treatment * Tmin	7	972.35	10.56	0.00	-478.60
	SVL + Osmo ini + timing + treatment + water access	8	973.31	11.52	0.00	-477.90
	SVL + Osmo ini + timing + treatment + Tmin	8	973.35	11.56	0.00	-477.92

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
		Fabien Aubret
	SVL + Osmo ini + timing + treatment + Tmax + treatment * Tmax	9	973.44	11.65	0.00	-476.77
	Osmo ini + timing + treatment + Tmin + treatment * Tmin	8	973.73	11.94	0.00	-478.11
	SVL + Osmo ini + treatment + Tmin + treatment * Tmin	8	974.33	12.54	0.00	-478.42
	SVL + Osmo ini + timing + treatment + Tmin + treatment * Tmin	9	975.72	13.93	0.00	-477.91
	Osmo ini		4	996.64	34.85	0.00	-494.12
	SVL + Osmo ini	5	998.62	36.83	0.00	-494.01
	Osmo ini + timing	5	998.69	36.90	0.00	-494.04
	SVL + Osmo ini + timing	6	1000.73	38.94	0.00	-493.94
	Null	F	3	1026.90	65.11	0.00	-510.33
	SVL	o r	4	1029.04	67.26	0.00	-510.32
		P
		e e r
		R e v
		i e w

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert, Fabien Aubret

Table S5 .

 S5 AICc based model selection comparing a null model (intercept only) to models testing relationships between female reproductive performance and environmental conditions. Models included the treatment timing (number of days between laying date and the end of water restriction), altitude and temperature indexes (T min and T max ) treated as linear covariates, while and hydric treatment and water access were treated as fixed factors. Population was set as a random factor in all models to account for intra-population variation and nonindependence. In analyses on offspring traits (SVL and BM), models also included mother treatment timing as linear co-variate, the juvenile sexes as fixed factor and mother identity as additional random factor to account for sibling non-independence. Preliminary investigations showed that female SVL did not explain variation in reproductive traits except for incubation time so this parameter was discarded from analyses on laying date, RCM, hatching success and offspring morphology to simplify model averaging. Highlighted models (bolded characters) are those explaining the greatest variation in response variables.

	Mother-offspring
		F
		o r
			P
	Reproductive performance	Models	e e r		k	AICc	ΔAICc	w i	LogLike
	Laying date	treatment + altitude + treatment * altitude altitude treatment + altitude R e v 6 4 5	669.06 669.54 671.03	0.00 0.48 1.97	0.46 0.36 0.17	-328.11 -330.57 -330.21
		treatment + Tmax + treatment * Tmax Tmax	6 4 i 681.78 684.99 e w	12.72 15.93	0.00 0.00	-334.46 -338.30
		Tmin			4	685.02	15.96	0.00	-338.31
		Null			3	685.21	16.15	0.00	-339.49
		treatment + Tmin + treatment * Tmin		6	685.72	16.66	0.00	-336.44
		treatment + Tmin			5	686.44	17.38	0.00	-337.92
		water access			4	686.45	17.39	0.00	-339.03
		treatment + Tmax			5	686.49	17.43	0.00	-337.94
		treatment			4	686.70	17.64	0.00	-339.15
		treatment + water access		5	687.99	18.93	0.00	-338.69
		treatment + water access + treatment * water access	6	690.15	21.09	0.00	-338.65

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
		Fabien Aubret
	Incubation time SVL + timing + altitude	6	242.90	0.00	0.30	-114.86
	SVL + timing + treatment + altitude	7	243.40	0.50	0.23	-113.90
	SVL + timing + treatment + altitude + treatment * altitude	8	245.31	2.41	0.09	-113.61
	SVL + altitude	5	246.06	3.16	0.06	-117.61
	SVL + timing + treatment	6	246.82	3.92	0.04	-116.82
	SVL + timing	5	247.66	4.75	0.03	-118.41
	SVL + timing + treatment + Tmax	7	247.73	4.82	0.03	-116.06
	timing + altitude	5	247.81	4.90	0.03	-118.49
	altitude	F	4	248.16	5.25	0.02	-119.80
	SVL + treatment + altitude o r	6	248.16	5.26	0.02	-117.49
	SVL + timing + Tmax P SVL + timing + treatment + Tmax + treatment * Tmax timing + treatment + altitude e e r	6 8 6	248.20 248.51 248.85	5.30 5.61 5.95	0.02 0.02 0.02	-117.51 -115.21 -117.83
	SVL + timing + treatment + water access SVL + timing + treatment + Tmin SVL + treatment + altitude + treatment * R e v 7 7 7 altitude SVL + timing + water access 6 SVL + timing + Tmin 6 i 249.09 249.24 249.30 249.93 249.96 e w	6.19 6.33 6.39 7.03 7.06	0.01 0.01 0.01 0.01 0.01	-116.75 -116.82 -116.85 -118.37 -118.39
	treatment + altitude	5	250.15	7.25	0.01	-119.66
	SVL + timing + treatment + Tmin + treatment * Tmin	8	250.44	7.54	0.01	-116.18
	SVL + timing + treatment + water access + treatment * water access	8	250.81	7.90	0.01	-116.36
	timing + treatment + altitude + treatment * altitude	7	251.01	8.10	0.01	-117.70
	treatment + altitude + treatment * altitude	6	251.79	8.89	0.00	-119.31
	timing		4	254.67	11.77	0.00	-123.06
	timing + treatment	5	254.85	11.95	0.00	-122.01
	SVL		4	255.00	12.10	0.00	-123.23

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
	Fabien Aubret
	SVL + Tmax	5	255.45	12.54	0.00	-122.31
	timing + Tmax	5	256.16	13.26	0.00	-122.66
	timing + treatment + Tmax	6	256.47	13.57	0.00	-121.64
	SVL + treatment + Tmax + treatment * Tmax	7	256.75	13.84	0.00	-120.57
	SVL + treatment	5	256.81	13.90	0.00	-122.99
	timing + Tmin	5	256.88	13.98	0.00	-123.02
	Null	3	256.93	14.03	0.00	-125.31
	timing + water access	5	256.96	14.05	0.00	-123.06
	SVL + Tmin F	5	257.00	14.09	0.00	-123.08
	timing + treatment + Tmin timing + treatment + water access o r	6 6	257.19 257.20	14.28 14.30	0.00 0.00	-122.00 -122.01
	SVL + water access P SVL + treatment + Tmax timing + treatment + Tmax + treatment * Tmax 7 5 6 e e r	257.26 257.33 257.70	14.36 14.42 14.80	0.00 0.00 0.00	-123.21 -122.07 -121.05
	Tmax treatment SVL + treatment + Tmin + treatment * Tmin R e v 4 4 7 SVL + treatment + Tmin 6 Tmin 4 i 258.04 258.65 258.66 258.91 258.97 e w	15.14 15.75 15.76 16.00 16.06	0.00 0.00 0.00 0.00 0.00	-124.75 -125.05 -121.53 -122.86 -125.21
	timing + treatment + Tmin + treatment * Tmin 7	259.10	16.19	0.00	-121.75
	water access	4	259.13	16.22	0.00	-125.29
	SVL + treatment + water access	6	259.14	16.23	0.00	-122.98
	timing + treatment + water access + treatment * water access	7	259.57	16.67	0.00	-121.98
	treatment + Tmax	5	259.81	16.91	0.00	-124.49
	treatment + Tmax + treatment * Tmax	6	259.98	17.07	0.00	-123.40
	treatment + Tmin	5	260.78	17.87	0.00	-124.97
	treatment + water access	5	260.91	18.01	0.00	-125.04

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
			Fabien Aubret
		SVL + treatment + water access + treatment * water access	7	261.28	18.38	0.00	-122.84
		treatment + Tmin + treatment * Tmin		6	261.78	18.87	0.00	-124.30
		treatment + water access + treatment * water access	6	263.24	20.33	0.00	-125.03
	RCM	timing + water access		5	66.14	0.00	0.22	-27.77
		timing			4	66.33	0.19	0.20	-28.97
		timing + Tmin			5	67.99	1.85	0.09	-28.70
		timing + treatment + water access + treatment * water access	7	68.20	2.06	0.08	-26.53
		timing + Tmax F			5	68.21	2.07	0.08	-28.80
		timing + treatment + water access timing + treatment o r		6 5	68.27 68.43	2.13 2.29	0.07 0.07	-27.71 -28.92
		timing + altitude	P		5	68.44	2.30	0.07	-28.92
		timing + treatment + Tmax + treatment * Tmax 7 timing + treatment + Tmin 6 e e r	69.52 70.07	3.37 3.93	0.04 0.03	-27.19 -28.61
		timing + treatment + Tmax timing + treatment + altitude timing + treatment + Tmin + treatment * Tmin 7 6 6 R e v	70.37 70.58 71.93	4.23 4.44 5.79	0.03 0.02 0.01	-28.76 -28.87 -28.39
		timing + treatment + altitude + treatment * altitude altitude	7 4 i e 72.85 88.13 w	6.71 21.99	0.01 0.00	-28.85 -39.87
		Tmax			4	89.12	22.98	0.00	-40.36
		treatment + altitude		5	90.33	24.19	0.00	-39.86
		treatment + Tmax			5	91.28	25.14	0.00	-40.34
		treatment + altitude + treatment * altitude		6	91.61	25.47	0.00	-39.38
		Null			3	92.27	26.13	0.00	-43.02
		treatment + Tmax + treatment * Tmax		6	93.33	27.19	0.00	-40.24
		Tmin			4	93.96	27.81	0.00	-42.78
		water access			4	94.35	28.21	0.00	-42.98
		treatment			4	94.41	28.27	0.00	-43.01

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

	Mother-offspring
	Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,
		Fabien Aubret
		treatment + Tmin		5	96.13	29.98	0.00	-42.76
		treatment + water access + treatment * water access	6	96.18	30.03	0.00	-41.66
		treatment + water access		5	96.53	30.39	0.00	-42.97
		treatment + Tmin + treatment * Tmin		6	98.37	32.22	0.00	-42.76
	Hatching success	timing + treatment + altitude + treatment * altitude	6	551.77	0.00	0.95	-269.47
		treatment + altitude + treatment * altitude		5	559.10	7.33	0.02	-274.25
		timing + treatment + Tmin + treatment * Tmin 6	559.80	8.02	0.02	-273.48
		timing + treatment timing F timing + treatment + altitude o r		4 3 5	565.66 567.34 567.48	13.89 15.57 15.71	0.00 0.00 0.00	-278.63 -280.55 -278.44
		timing + treatment + Tmax P		5	567.79	16.02	0.00	-278.60
		timing + treatment + Tmin timing + treatment + water access e e r timing + treatment + water access + treatment * water access treatment + Tmin + treatment * Tmin timing + altitude R e v 5 5 6 5 4	567.80 567.82 568.04 569.14 569.19	16.03 16.04 16.27 17.37 17.41	0.00 0.00 0.00 0.00 0.00	-278.60 -278.61 -277.60 -279.27 -280.40
		timing + Tmax timing + water access timing + Tmin	4 4 4 i 569.38 569.40 569.49 e w	17.60 17.63 17.71	0.00 0.00 0.00	-280.49 -280.51 -280.55
		timing + treatment + Tmax + treatment * Tmax 6	569.56	17.79	0.00	-278.36
		treatment + altitude		4	570.61	18.84	0.00	-281.11
		altitude		3	571.80	20.03	0.00	-282.78
		treatment		3	572.54	20.77	0.00	-283.15
		Null		2	573.19	21.42	0.00	-284.54
		treatment + Tmax		4	573.38	21.61	0.00	-282.49
		treatment + Tmax + treatment * Tmax		5	573.60	21.82	0.00	-281.50
		Tmax		3	573.90	22.13	0.00	-283.84

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert, Fabien Aubret

	Mother-offspring
	treatment + Tmin		4	574.47	22.70	0.00	-283.04
	treatment + water access		4	574.63	22.85	0.00	-283.12
	treatment + water access + treatment * water access	5	574.94	23.17	0.00	-282.17
	Tmin		3	575.03	23.26	0.00	-284.40
	water access		3	575.29	23.51	0.00	-284.53
	Offspring SVL sex + altitude		6	960.54	0.00	0.29	-474.14
	sex + treatment + altitude		7	962.07	1.53	0.14	-473.86
	sex + treatment + altitude + treatment * altitude 8	962.08	1.55	0.13	-472.82
	timing + sex + altitude F		7	962.42	1.88	0.11	-474.04
	treatment + sex + altitude + sex * altitude o r		8	963.96	3.42	0.05	-473.76
	timing + sex + treatment + altitude timing + sex + treatment + altitude + treatment * altitude P sex + Tmax e e r timing + treatment + treatment + altitude + sex * altitude sex timing + sex + Tmax R e v 8 9 6 9 5 7	964.10 964.19 965.04 966.00 966.23 966.66	3.56 3.66 4.50 5.46 5.69 6.12	0.05 0.05 0.03 0.02 0.02 0.01	-473.83 -472.82 -476.39 -473.72 -478.02 -476.16
	sex + treatment + Tmax sex + treatment + sex * treatment timing + sex	7 7 6 i 966.86 967.40 967.80 e w	6.32 6.87 7.26	0.01 0.01 0.01	-476.26 -476.53 -477.77
	sex + treatment		6	968.05	7.51	0.01	-477.89
	sex + Tmin		6	968.12	7.58	0.01	-477.93
	sex + water access		6	968.14	7.60	0.01	-477.94
	sex + treatment + Tmax + treatment * Tmax		8	968.27	7.74	0.01	-475.91
	timing + sex + treatment + Tmax		8	968.63	8.10	0.01	-476.09
	treatment + sex + Tmax + sex * Tmax		8	968.82	8.28	0.00	-476.19
	timing + sex + treatment + sex * treatment		8	969.09	8.56	0.00	-476.32
	timing + sex + water access		7	969.64	9.11	0.00	-477.65

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,

	Mother-offspring
	Fabien Aubret
	timing + sex + Tmin	7	969.73	9.19	0.00	-477.69
	timing + sex + treatment	7	969.78	9.24	0.00	-477.71
	sex + treatment + Tmin	7	969.92	9.39	0.00	-477.79
	sex + treatment + water access	7	969.96	9.43	0.00	-477.81
	treatment + sex + water access + sex * water access	8	970.03	9.49	0.00	-476.79
	timing + sex + treatment + Tmax + treatment * Tmax	9	970.21	9.67	0.00	-475.83
	timing + treatment + sex + Tmax + sex * Tmax 9	970.61	10.08	0.00	-476.03
	timing + treatment + sex + water access + sex * water access F	9	971.60	11.06	0.00	-476.52
	timing + sex + treatment + water access timing + sex + treatment + Tmin o r	8 8	971.64 971.70	11.10 11.16	0.00 0.00	-477.60 -477.63
	treatment + sex + Tmin + sex * Tmin P	8	971.82	11.28	0.00	-477.69
	sex + treatment + water access + treatment * water access sex + treatment + Tmin + treatment * Tmin e e r	8 8	971.98 972.02	11.45 11.49	0.00 0.00	-477.77 -477.79
	timing + treatment + treatment + Tmin + sex * Tmin timing + sex + treatment + water access + treatment * water access R e v 9 9	973.62 973.66	13.08 13.12	0.00 0.00	-477.53 -477.55
	timing + sex + treatment + Tmin + treatment * Tmin altitude i 9 973.78 5 992.06 e w	13.24 31.53	0.00 0.00	-477.61 -490.94
	timing + altitude	6	993.84	33.30	0.00	-490.79
	Tmax	5	997.55	37.01	0.00	-493.68
	Null	4	998.25	37.71	0.00	-495.06
	timing + Tmax	6	999.06	38.52	0.00	-493.40
	timing	5	999.69	39.15	0.00	-494.75
	treatment	5 1000.08	39.54	0.00	-494.95
	Tmin	5 1000.12	39.58	0.00	-494.97
	water access	5 1000.18	39.64	0.00	-495.00
	timing + water access	6 1001.57	41.03	0.00	-494.66

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert,

	Mother-offspring
		Fabien Aubret
	timing + Tmin		6 1001.61	41.07	0.00	-494.67
	timing + treatment		6 1001.68	41.15	0.00	-494.71
	Offspring BM altitude		5 3062.26	0.00	0.21	-1526.04
	sex + treatment + sex * treatment	7 3062.87	0.61	0.15	-1524.26
	sex + altitude		6 3063.27	1.01	0.13	-1525.51
	timing + altitude		6 3064.27	2.01	0.08	-1526.01
	sex + treatment + altitude		7 3064.90	2.64	0.06	-1525.28
	timing + sex + treatment + sex * treatment	8 3064.97	2.71	0.05	-1524.26
	timing + sex + altitude F		7 3065.30	3.04	0.05	-1525.48
	Tmax sex + treatment + altitude + treatment * altitude 8 3066.77 5 3065.81 o r	3.54 4.51	0.04 0.02	-1527.81 -1525.16
	timing + sex + treatment + altitude P	8 3066.79	4.53	0.02	-1525.17
	treatment + sex + altitude + sex * altitude sex + Tmax e e r	8 3066.93 6 3066.94	4.66 4.68	0.02 0.02	-1525.24 -1527.34
	Null timing + Tmax sex	4 3067.16 6 3067.88 5 3068.22 v R e	4.89 5.62 5.95	0.02 0.01 0.01	-1529.52 -1527.81 -1529.02
	timing + sex + treatment + altitude + treatment * altitude water access i 9 3068.56 5 3068.72 e w	6.29 6.45	0.01 0.01	-1525.00 -1529.27
	sex + treatment + Tmax		7 3068.77	6.51	0.01	-1527.21
	treatment + sex + Tmax + sex * Tmax	8 3068.82	6.55	0.01	-1526.19
	timing + treatment + sex + altitude + sex * altitude	9 3068.83	6.57	0.01	-1525.14
	treatment		5 3068.97	6.70	0.01	-1529.39
	timing + sex + Tmax		7 3069.03	6.77	0.01	-1527.34
	Tmin		5 3069.12	6.85	0.01	-1529.47
	timing		5 3069.22	6.95	0.01	-1529.52
	sex + water access		6 3069.80	7.54	0.00	-1528.77
	sex + treatment		6 3070.04	7.78	0.00	-1528.89

conflict for water varies across altitude but is mitigated in the oviparous form of the bimodal lizard Zootoca vivipara

  Andréaz Dupoué, Mahaut Sorlin, Murielle Richard, Jean-François Le Galliard, Olivier Lourdais, Jean Clobert, Fabien Aubret

Acknowledgments

449

We thank Pierre-Louis Duarte and Loïse Bessaguet for their help on the field and Pauline 450 Blaimont for English revising and helping during experiments. We are also grateful to the

Data availability 464

Data will be made available should the manuscript be accepted. 465

Authors' contribution

466

AD conceived the study, captured individuals, led the experiments, performed the analyses, 467 and draft the manuscript. MS and MR helped in capture and performed the experiments.