Posted on Authorea 9 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161526176.66765405/v1 — This a preprint and has not been peer reviewed. 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All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161526176.66765405/v1 — This a preprint and has not been peer reviewed. 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Open Soc. organisms, other Alstr in found parti- is future. pattern be in same would clades the References It or whether dramatically. species test niches widespread to their strategies such framework expand in life same related the to especially assessments use or clades innovations to further particular interesting morphological exist, these cularly key allowed do of have innovations mechanisms that morphological molecular combined cope key understand by to to and seek accompanied toads size should helping shifts body by niche enlarged success true as expansion Recognizing such increase Factors to stability. shown conditions. and novel been unfilling innovations with have niche morphological performance key than locomotor accompanies climates enhanced novel indeed expansion in presents niche unfilling preadaption niche by of while caused clades, divergence C E. niche and Meanwhile, B E. clade. A, b E. expansion W. in niche in unfilling niche just realized than prevalent of more degree are expansion the morpho- in within and variations clades models of considerable the niche shown across correlative have unfilling with we and areas geographic models, different evolution from logical clades genetic studies. distinct further Combing in will needed trade-offs and are resulting length adapting models the optimization body towards of Conclusion and between consequences expansion tools fitness variation niche analytical the underlie to novel quantifying that challenging, in contribute complex, mechanisms be endurance be together proximate can and may the performance speed HLL SVL) locomotor because LAHL, for (i.e. (i.e Indeed, SVL plasticity changes niche. phenotypic seems enlarged evolution found novel there combined we and Collectively, exist, performance 2003). 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Supplementary different character A, BM clades, these completely E. fitted Interestingly, remained clade best trend. in the same signal, 2010, mm the in phylogenetic al. the have 27.19 mm et with value: FL Phillips 21.77–23.57 the and (mean accord 2014, C, HLL LAHL overcoming in al. study, E. helps are et our in which that (Tingley In 23.95mm habitats, studies toads 2010). novel prior al. in by in et preadaptation highlighted Kolbe temperature key model high mechanistic a clades. or shifts C be evolutionary precipitation E. important may attributes insufficient and traits morphological B shifted of have These to E. challenge newly tends A, performance. niche in E. shifted locomotor release) newly in of of for SVL (ecological evolution tolerance enlarged Yoder species competitors the for found 1997, influences reduced we reason Queiroz Moreover, temperature highlight likely and body most results (Losos preferred the Our SVL of be 2011). larger maintenance may al. possi- niche that for second likely allow et The is may (Oufiero 1982). third areas SVL (Mautz The shifted SVL 2010). novel and al. in in endurance et release variation fasting ecological between SVL is relationship of bility expected pattern the observed to the pertains underlie may factors Several .boulengeri S. m .e l 05 rmtcncesit n opooia hnei w nua idseis R. – species. bird insular two in change morphological and shifts niche Dramatic 2015. al. et P. ¨ om, .–Po N 8:e34.Bobr,S .e l 03 etn o hlgntcsga in signal phylogenetic for Testing 2003. al. et P. S. Blomberg, e43641. (8): 7 ONE PLoS – ). od,ncedvrec cusacmaidb ih xaso ahrta nlig niche unfilling, than rather expansion niche by accompanied occurs divergence niche toads, .boulengeri S. od nteQnhiTbtPaeurgo.I h case the In region. Plateau Qinghai-Tibet the in toads 8 .boulengeri S. lds n possibility One clades. Perca Posted on Authorea 9 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161526176.66765405/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. eprtr atrscmie ihhmniflecsrdsg h naience lb hneBo.19: Biol. Change Glob. – niche. invasive World the Old redesign influences of human 3406–3422. Evol. Invasion with Ecol. 2013. combined Trends al. patterns – temperature et invasions. biological Y. from W. Insights Guo, studies: shift niche Unifying speciation 260–269. 2014. explore 29: al. to A. et models 1781–1793. A. Grafen, niche 58: environmental Guisan, 315–325. 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A. Gans, al. Mulch, – et In monsoon. P. – Indian Molnar, loss. the 443–481. water and pp. evaporative Plateau Press, of Academic Patterns reptilian. 1982. the J. W. Mautz, > iuu hartii Rivulus . .–Eooy9:170–179. 92: Ecology – ). 11 < tp:/ihbcm/liam- https://github.com hnlamarina Rhinella .–Mol. – ). Posted on Authorea 9 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161526176.66765405/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. aa Q,CSadF eddtewiig l uhr otiue iheisadcmet oadthe toward the comments analyzed and CKS edits and with XQL contributed data; authors raw All the writing. processed the FX leaded and manuscript. FX XQL final and JPJ; CKS and XQL, JHH data; MHZ, XXS, YMH, BON). from (Sino Networks contributions Science Observation Ecology Sichuan Biodiversity Author of of China Ministry Platform and Conditions the 2019JDPT0020 Basic Department of of Technology Project Construction and Assessment 2019HJ2096001006, China and Environment, Survey work. and Professor Biodiversity field and the in Shu help 2019QZKK05010503), Guocheng (STEP, kind Dr. Qi, their Yin for Dr. Xu – Li, Funding Xiancheng Cheng and Dr. Shi thank We Shengchao draft. Gong, earlier Dajie the of conception on help molecular streamlined 348–355. for Acknowledgements 20: Resour. platform Ecol. at desktop Mol. Appendix scalable – material and studies. Supplementary phylogenetics integrated evolutionary an and PhyloSuite: management 2020. data al. sequence 23: 45–53. et Biol. 121: (Lepidosauria, D. 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D. of – C. accuracy India. Thomas, the Himalaya, Measuring Toad 1988. in Lazy lake J. Boulenger’s altitude Swets, of high 7655–7663. record from first heights: Megophryidae) new Anura: Scaling bia: 2015. al. et B. Subba, hssuywsfne yteScn iea lta cetfi xeiinadRsac Program Research and Expedition Scientific Plateau Tibetan Second the by funded was study This etakCuln i rmFeeUniversit Freie from Liu Chunlong thank We – Q n Xcnevdteie;XLadF olce h aawt assistance with data the collected FX and XQL idea; the conceived FX and XQL – < www.oikosoffice.lu.se/appendix 12 tBri nvri´ ai alyfor Saclay Universit´e Paris & Berlin ¨ at > . ctgrboulengeri Scutiger (Amphi- Posted on Authorea 9 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161526176.66765405/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. pce yclsi h aierne h oi n ahdcnorlnsilsrt,rsetvl,10 and 100% respectively, components. illustrate, principal five lines first contour the the of dashed on range. occurrences and Loadings native of 1 solid while the density Table The range, in the non-native environment range. indicates available and native Shading the native the range. of the occu- shifted in 50% both only the in cells climates in occupied represent unfilling by expansion climates areas and species calibrated niche indicate overlap Green PCA, blue indicate expansion, clade. a and Niche areas each range of lines). red for native axes solid space the two (red environmental in first China the pied the in in available by stacked conditions depicted are of space situations range environmental entire in the ranges on ( shifted ( bi-directions; and values; native in observed ween distributions represent similarity lines niche diamond distributions, of equivalency niche of Histograms geri within clades (a): edge. each 3. along Figure states the interpolating elevation ( and bold; PCs in tree dated (c): in of clades River; (Ma) six time Min of divergence MR: range suitable River; the Dadu and DDR: River; 2. 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All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161526176.66765405/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. aibe htdces h antems hnte r mte r ie o ahclade. each clades. invaded for and * the given range clade; native are each between omitted for indices are bold changes in they Niche given when are 3 most Table isolation the in gain used hinge. the when = decrease gain H highest that with variables quadratic, variables = environmental Q The linear, 2. = L Abbreviations: 1. Note: model. Maxent in variables environmental the of contributions relative corresponds and mainly parameters PC2 while The temperatures, 2 Table to are corresponds three mainly first PC1 occupied Loadings component. precipitation. each 1. to than for greater bold eigenvalues with in components given principal all retained We Note: ld aaee U i1bo i3bo i8bo2bo4bio15 bio14 bio12 bio8 bio7 b W. bio3 a W. bio2 D E. C E. bio1 B E. AUC A E. Parameter Clade lvto 094-.5 .7 .3 -0.048 0.232 0.383 0.274 -0.142 -0.055 -0.278 -0.914 -0.123 0.199 PC5 PC4 PC3 PC2 (%) variance PC1 of percentage Cumulative (%) variance of Percentage Eigenvalue Elevation Dissimilarity quarter coldest of Precipitation quarter warmest of Precipitation quarter driest of Precipitation quarter wettest of Precipitation seasonality Precipitation month driest of Precipitation precipitation Annual Variables rcptto fwtetmonth wettest of Precipitation Q .7 .1 .1 .2 0 0.026 0 0 0.062 0 0 0.244 0.914 0.892 0.313 0.194 0.99 0.012 0.937 LQH4 0.977 L2.5 0.979 LQ3 LQ2 LQ1 LQ1 .AW .7 .2 .1 0.285 0.715 0.683 0.017 0.822 0.070 0.782 0.459 0.782 0.065 0.002 0.792 b 0.842 0.002 W. b 0.812 0.095 0.485 W. a 0.011 0.743 W. B E. a 0.940 0.001 W. A E. U a 0.110 0.980 W. D E. a 0.881 0.220 W. C E. D 0.252 E. B S E. D E. A E. D E. C E. C E E. B E. C test E. A Sim. E. B E. B E. A E. D A E. clades Native Clades (bio12) (bio15) 0.652* 0.231 (bio14) (bio17) (bio13) (bio19) (bio16) (bio18) .8 .1 .4 0 0.049 0.115 0 0.153 0.301* 0 0.163 0.184 0 0 0 0.12 0.033 14 0.010 0.703 0.373 0.368 9635.7 0188.2 91.138 1.516 7.217 83.921 2.882 13.723 70.198 17.322 52.876 3.638 23.253 29.623 4.883 29.623 0.097 6.221 -0.500 0.431 -0.231 -0.770 0.347 -0.224 0.206 -0.216 0.353 -0.317 -0.624 -0.080 0.241 -0.012 -0.361 013082-.4 0170.051 -0.213 -0.137 -0.348 0.872 -0.176 -0.173 .2 .2 .7*0.002 0 0.174* 0.119 0.254* 0.001 0 0.174 0.026 0 0.255* 0.029 0.006 0.01 0.001 0.917 0.935 0.939 0.913 0.159 0.894 0.986 0.990 0.907 0.956 0.997 0.700 0.610 .5 0160.217 -0.166 0.153 0.879 .2 0140.141 -0.124 0.127 0.135 -0.033 0.191 .8 0.899 0.087 0.838 0.841 0.970 0.106 0.979 0.014 0.656 0.010 0.744 0.093 0.986 0.044 0.611 0.003 0.300 0.603 0.515 0.390 0.097 .3 -0.425 0.634 0.266 -0.176 0.752 0.289 0.302 .0*0.172 0.308* Posted on Authorea 9 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161526176.66765405/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. Elev. PC5 PC4 PC3 PC2 PC1 oe brvain:K lmegsK M rwinmto;E:Erybrt N ht os;OU: noise; White Akaike corrected WN: AICc: burst; logarithm; Early Neperian EB: for values motion; L: ln Brownian - BM: peak; adaptative K; one Blomberg’s with Ornstein-Uhlenbeck K: Abbreviations: Note: FL TFL TW TL HLL LAD LAHL ED UEW IOS INS SL HW HL SVL characteristic size-corrected on based analyses fitting OU: model and noise; test signal data. White phylogenetic Akaike of corrected WN: results The AICc: burst; 5 logarithm; Table Early Neperian EB: for values motion; L: ln Brownian criterion; - BM: information peak; adaptative K; one Blomberg’s with Ornstein-Uhlenbeck K: Abbreviations: Note: and components principal climate five for obtained significant analyses indicate fitting values model Boldface and of Elevation. signal p-value unfilling. Phylogenetic the U: 4 test: stability; Table S: Sim. clades; expansion; pairwise niche p between E: with value similarity test; overlap the similarity D: niche Schoener’s the D: Abbreviations: Note: .6 (0.045) 1.568 (0.049) 1.381 (0.04) 1.648 .5 02)-.4 748048-.1 748003-.4 7480.003 0.002 37.488 42.066 -12.033 -9.744 0.002 0.003 42.017 37.438 -12.009 -9.719 0.353 0.428 32.066 27.488 -12.033 -9.744 (0.09) 1.141 (0.33) 0.749 (0.21) 0.950 P-vle lLAICc -lnL value) - (P K Pvle MO BWN EB OU BM (P-value) K .7 09)846-.7 .9 .9 .0 .0 .8 .2 0.001 0.002 0.002 1.027 -14.476 -7.964 16.238 8.486 12.982 0.001 0.002 0.001 0.002 0.002 -21.951 -14.648 -9.616 0.605 19.975 -8.218 16.324 0.001 13.808 0.001 13.109 0.002 -22.364 8.697 0.288 0.002 -22.359 0.002 -15.901 20.182 0.243 0.092 -24.470 20.180 0.002 -26.583 16.951 0.001 -9.798 -17.964 0.002 -32.365 22.291 0.002 -22.610 13.899 16.235 -8.973 0.096 -25.914 25.182 0.002 -16.026 20.305 12.982 0.297 (0.65) -31.951 0.587 21.957 0.002 -26.819 17.013 8.486 0.189 (0.67) -19.616 0.565 0.003 -32.547 22.409 0.003 19.975 0.296 (0.93) -32.364 0.376 -26.098 25.274 0.003 -16.646 13.808 0.310 (0.96) -25.850 0.359 22.049 -17.149 17.323 20.182 0.314 (0.66) -36.583 0.565 17.575 0.003 16.925 0.357 (0.75) -42.365 0.502 0.003 -16.821 22.291 (0.35) -35.914 0.804 -17.296 17.410 25.182 (0.37) 0.749 17.648 21.957 0.410 (0.57) 0.606 0.443 (0.21) -26.646 0.846 (0.09) -27.149 1.142 17.323 (0.08) 1.109 17.575 (0.18) 0.831 (0.12) 1.066 (0.07) 1.222 ldsNtv ldsDSm etESU S E test Sim. D clades Native Clades .aW 0.527 0.444 0.307 b W. b W. b W. a W. D E. C E. < .5adtepeerdncesitmodel. shift niche preferred the and 0.05 ω: kiewihs lv lvto.Blfc ausidct h rfre model. preferred the indicate values Boldface Elevation. = Elev. weights. Akaike lLAICc -lnL 446-0920.546 0.507 -40.952 -39.837 24.476 23.918 0.617 -9.760 8.880 4.69.2 0.687 0.572 98.121 0.665 17.716 -45.06 26.532 -4.858 -9.266 9712.6 0.627 27.561 -9.781 MO BWN EB OU BM ω ω 15 0.010 0.020 0.050 4882.1 .0 4872.9 .0 5161.3 0.420 0.304 0.324 18.332 99.754 27.972 -45.877 -5.166 0.005 -9.986 0.004 108.121 0.005 27.693 -45.06 36.523 -4.847 0.005 -9.262 0.004 108.121 0.006 27.716 -45.06 35.881 -4.858 -8.941 9703.5 .0 9713.6 .0 1.2 8680.364 28.648 -10.324 0.004 37.561 -9.781 0.004 37.559 -9.780 lLAICc -lnL 455-1000042.7 3.5 .0 426-0510.447 0.375 0.486 -40.551 -39.751 24.276 -8.764 23.876 0.004 8.382 0.003 -30.952 0.004 -29.837 24.476 23.918 0.004 0.240 0.004 -31.030 8.880 -29.943 24.515 0.004 23.971 0.228 AICc 8.886 -lnL 0.031 0.037 0.304 ω ω 0.969 0.963 0.696 lLAICc -lnL lLAICc -lnL 0.514 0.628 0.493 ω ω lLAICc -lnL 712-6250.708 0.754 -26.265 0.906 -20.231 17.132 0.903 -13.545 14.115 -36.439 10.773 0.698 22.220 0.808 -21.324 0.700 -35.268 14.662 0.685 -27.574 21.634 0.682 -38.165 17.787 0.638 -43.918 23.083 -37.071 25.959 22.536 0.585 0.551 -27.357 -27.583 17.678 17.791 1.3 0880.642 0.566 30.868 26.930 -11.434 -9.465 lLAICc -lnL ω ω Posted on Authorea 9 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161526176.66765405/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. C+C 66 .953 331.91.11 .81. 57 08 94 26 25.83 22.06 -2.65 23.5 24.32 -8.24 -9.41 -3.44 22.35 -8.94 25.46 1.79 -0.82 26.01 -8.46 23.53 -4.47 -1.05 -4.49 -5.73 -2.66 -3.47 26.12 -1.8 -12.06 -5.15 -3.96 25.2 2.56 -7.58 -13.83 -7.8 3 -3.84 -11.53 41.64 0.7 -0.61 -1.08 -3.02 -5.48 -10.51 -3.88 -4.68 1.02 17.1 -5.14 17.97 -11.83 -5.09 -6.28 -134.43 -5.3 -4.39 -3.83 -2.73 -3 -27.97 17.07 -135.34 -3.19 6.28 3.98 -3.13 -9 -30.87 0.88 -28.04 -34.46 -130.89 -32.64 -5.52 3.02 16.58 16.64 4.73 -33.95 -108.99 -5.86 -0.43 12.4 15.88 -33.82 -27.43 -37.43 12 -127.08 18.57 11.39 1.73 -124.89 12.07 -30.37 6.92 -27.23 -30.22 -30.66 18.28 18.73 -105.5 0.88 19.01 -109.16 12.23 6.9 13.16 4.39 -34.98 18.75 -121.38 12.52 -26.85 -35.03 -29.8 10.09 -101.8 16.76 10.39 1.01 19.15 -108.14 12.37 -28.66 6.11 -11.41 12.8 -26.49 -115.07 42.38 18.75 9.96 -29.11 43.3 -101.3 -11.71 6.04 13.5 19.44 9.86 -18.54 -12.91 -78.5 42.76 12.47 18.97 73.99 -112.79 -2.41 -23.03 -13 42.24 12.46 18.72 -17.13 10.41 -22.37 -144.88 -78.57 5.37 41.64 19.41 9.02 -16.36 4.66 -17.65 42.97 -10.73 19.38 -143.58 -141.71 -16.8 3.28 12.05 9.83 10.16 44.08 -11.25 -17.31 1.69 8.95 48.33 10.15 42.67 -149.04 -10.55 42.38 1.06 -74.42 5.36 -143.73 65.24 -19.76 -10.25 1.21 14.11 6.36 43.32 66.64 -17.82 3.33 -83.85 12.84 PC1+PC2+PC3+PC4 64.83 -18.93 4.38 4.61 -0.39 -84.8 13.63 PC1+PC2+PC3+PC5 -17.46 77.93 -6.75 13.55 PC1+PC2+PC4+PC5 1.84 -20.55 73.99 -14.08 PC1+PC3+PC4+PC5 -24.63 66.98 -26.09 -7.3 PC2+PC3+PC4+PC5 -17.77 70.01 -25.48 -4.16 PC4+PC5 65.82 -27.12 PC3+PC5 48.33 66.02 -26.64 PC3+PC4 44.26 65.89 PC2+PC5 37.16 PC2+PC4 48.42 PC2+PC3 PC1+PC5 PC1+PC4 PC1+PC3 PC1+PC2 PC5 PC4 PC3 PC2 C 05 3.1-36 44 1.9-05 1.6-35 1.2-63 3.6-32.73 -34.46 -26.39 -11.42 -23.58 -16.26 -10.57 -18.89 14.46 -23.63 -32.91 40.54 PC1 scutiger-boulengeri-anura-megophryidae climatic-niche-evolution-with-key-morphological-innovations-across-clades-within- legends.pdf and p Figures with difference Significant * file model; Hosted preferred the indicate values Boldface Note: traits. and components principal climate Model between value AICc on based comparison Model 6 Table criterion; information ω: kiewihs odaevle niaetepeerdmodel. preferred the indicate values Boldface weights. Akaike 7.4-4.3-3 -78.14 -138 -147.83 -70.04 -159.79* -103.42* V A WS ALTLF LHLT N O E DTW ED UEW IOS INS TL HLL HL FL TFL LAHL SL HW LAD SVL vial at available https://authorea.com/users/400321/articles/512661- 16 130 7.6-1.2-0.3-0. 115-0.1-2.3-3.9-4.2-3.8-3.5-117.89 -138.25 -134.88 -142.72 -139.79 -129.03 -106.91 -121.5 -108.7 -101.33 -113.52 -78.8* -78.36 -151.84* -143.09 146 114 191 129 180 178 181 185 163 109 -118.63 -140.92 -136.33 -138.56 -138.13 -133.87 -127.89 -109.56 -108.09 -127.62 -122.99 -109.78 -102.34 -109.15 -116.42 -101.44 -114.63 < 0.05. AICc 134 123 -138.46 -132.34 -147.65 -133.48 -141.13 154-3.5-111.05 -138.52 -136.75 -135.4 -40.5* -149.98 -161.48* 3.2-3.61 -33.42 -148.49* -126.25