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Shoot Flammability of Vascular Plants Is Phylogenetically Conserved And

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Shoot Flammability of Vascular Plants Is Phylogenetically Conserved And DispatchDate: 30.03.2020 · ProofNo: 635, p.1 LETTERS https://doi.org/10.1038/s41477-020-0635-1 1 2 3 Shoot flammability of vascular plants is 4 5 6 phylogenetically conserved and related to habitat 7 8 fire-proneness and growth form 9 10 1 1 2 1 3 11 Xinglei Cui ✉ , Adrian M. Paterson , Sarah V. Wyse , Md Azharul Alam , Kévin J. L. Maurin , 12 Robin Pieper1, Josep Padullés Cubino1,4, Dean M. O’Connell1, Djessie Donkers1, Julien Bréda1, 13 5 6 1 14 Hannah L. Buckley , George L. W. Perry and Timothy J. Curran ✉ 15 16 . Q1 17 Terrestrial plants and fire mhave interacted for at least 420 mil- flammability is to evaluate variation in flammability with phyloge- 18 lion years. Whether recurrent fire drives plants to evolve netic approaches, but few such studies have been reported. These 19 higher flammability and what the evolutionary pattern of previous studies have mostly focused on specific genera20 and used .. 20 plant flammability is remain unclear. Here, we show that phy- qualitative, rather than quantitative, measures of flammability17,21. Q3 Q4 Q5 Q6 . mm Q2 21 logeny, fire-pronenessm of habitat and growth form are impor- We burned shoots (length, 70 cm) of 194 species (120 indig- 22 tant predictors of the shoot flammability of 194 indigenous enous to New Zealand and 74 exotic species introduced from 23 and introduced vascular plant species (Tracheophyta) from other parts of the world) from across the Tracheophyta (vascular . Q7 24 New Zealand. The phylogenetic signal of the flammability plants; Supplementary Fig. 1, mTable 1). We measured four compo- 25 components and the variation in flammability among phylo- nents of flammability: ignition frequency (ignitability), burning 26 genetic groups (families and higher taxonomic level clades) time (sustainability), maximum temperature (combustibility) and 27 demonstrate that shoot flammability is phylogenetically con- burnt biomass (consumability), and related these to phylogeny, the 28 served. Some closely related species, such as in Dracophyllum fire-proneness of the species’ habitat and growth form of the spe- 29 (Ericaceae), vary in flammability, indicating that flamma- cies. The selected species showed a wide range of shoot flamma- 30 bility exhibits evolutionary flexibility. Species in fire-prone bility attributes: 22 species did not ignite on our device (ignition 31 ecosystems tend to be more flammable than species from frequency of zero), whereas 82 species ignited in 100% of samples. 32 non-fire-prone ecosystems, suggesting that fire may have an Mean consumed biomass per species ranged from 0% to 94%, the 33 important role in the evolution of plant flammability. Growth mean maximum temperature per species reached 771.5 ± 23.0 °C 34 form also influenced flammability—forbs were less flamma- (mean ± 1 s.e.m.), and mean burning times ranged from 0 s to 35 ble than grasses, trees and shrubs; by contrast, grasses had 240 s (Supplementary Table 1). Combustibility, consumability and 36 higher biomass consumption by fire than other groups. The ignitability were strongly positively correlated, whereas sustain- 37 results show that shoot flammability of plants is largely cor- ability had a weaker correlation with ignitability and consumability 38 related with phylogenetic relatedness, and high flammability (Supplementary Fig. 2). 39 may result in parallel evolution driven by environmental fac- The integration of flammability data with the phylogeny showed 40 tors, such as fire regime. that closely related species tend to have similar flammability 41 Fire has affected the distribution and evolution of terres- (Fig. 1), although flammability varied considerably among some 42 trial plants for at least 420 million years1–6 and many species have 43 developed adaptations to persist in the face of this disturbance5,7. 44 Although a growing number of researchers support the idea that Q8 Table 1 | Results ofm the phylogenetic tests for flammability 45 fire has selected some plant species to become more flammable8–12 components across selected indigenous and introduced 46 or, in some cases, less flammable11,13, others have argued that New Zealand vascular plants species 47 flammability has not evolved in response to fire, but is the result 48 of exaptations, whereby traits fulfilling other functions also influ- Flammability components Pagel’s λ 49 ence flammability14–16. Although there is evidence in some taxa λ value P 50 that plant flammability has evolved in response to changes in fire Ignition frequency 0.74 <0.001 51 regimes10,17,18, broad-scale phylogenetic patterns in plant flamma- Burning time 0.27 0.005 52 bility remain unclear. A better understanding of the evolution of 53 flammability would facilitate our understanding of the long-term Maximum temperature 0.51 <0.001 54 interactions between fire and plants, and may help to prepare us for Burnt biomass 0.48 <0.001 55 a warmer world, in which fire risk may be higher in many regions19. P values give the significance of Pagel’s (n 190 species). 56 One method to decipher the evolutionary patterns of plant λ = 57 58 59 1Department of Pest-management and Conservation, Lincoln University, Lincoln, New Zealand. 2Bio-Protection Research Centre, Lincoln University, 60 Lincoln, New Zealand. 3School of Science, The University of Waikato, Hamilton, New Zealand. 4Department of Ecology, Evolution and Behavior, University 61 of Minnesota, St Paul, MN, USA. 5School of Science, Auckland University of Technology, Auckland, New Zealand. 6School of Environment, University of 62 Auckland, Auckland, New Zealand. ✉e-mail: [email protected]; [email protected] 63 A B NatURE Plants | www.nature.com/natureplants DispatchDate: 30.03.2020 · ProofNo: 635, p.2 LETTERS NATURE PLANTS APOsim CARwak CARcor CHImac ANTslo JUNgre LUZpum 64 LUZruf CROcro PHOten PHOcoo CORaus CHIrig RYTset OPHpla 65 BROhor POAcit AGApra POAcol 64.9 35.2 78.6 50.5 25.0 FESnov CLImin 25.0 DACglo 86.3 0.0 RIPsca ANTodo 66 0.0 AMMare 93.5 HAKser 23.5 15.0 DEYave 3.1 89.2 10.0 PROner 74.4 19.57 0.9 68.9 AGRmue 11.4 26.0 7.5 20.3 49.2 31.2 AGRcap 0.3 KNIexc 0.7 56.3 PIPexc 67 0.0 4.3 88.8 PRUspc 0.0 0.0 PSEcol 2.3 1.1 83.1 1.0 239.9 2.4 12.5 PRUlau 15.9 43.1 169.8 13.1 MAGgra 750.8 5.0 666.8 77.1 22.5 492.6 5.3 0.5 210.4 68 604.8 PRUser 16.7 0.0 HEDarb 150.0 2.4 4.4 433.5 150.0 26.9 239.5 78.6 43.3 PHOgla 0.0 8.6 LAUnob 246.6 0.3 BEItar 1.9 44.56 12.5 53.8 284.9 729.1 215.0 PYRcom 40.5 185.3 2.6 69 152.9 618.3 9.2 10.00 490.8 10.00 BEItaw 3.3 36.3 167.7 580.9 0.0 100.0 8 150.0 25.0 284.2 32.0 MALdom 7 43.8 25.4 330.1 87.5 0.0 32.3 GINbil 213.5 . 600.5 25.0 0.0 5 13.6 28.3 375.9 24.5 70 RUBcis 13.1 584.3 50.0 25.0 189.5 PODtot 100.0 2.7 87.5 25.0 510.8 1 0.00 27.9 41.4 3.7 12.5 100.0 207.1 100.0 9.2 RUBfru 525.1 100.0 83.3 12.4 PODhal 20.3 7.4 150.0 75.0 4.6 48.3 613.9 0.0 543.3 71 13.6 50.0 100.0 29.8 DACdac GEUlei 100.0 100.0 476.0 232.3 12.5 6.4 12.2 14.1 100.0 279.8 25.0 5.8 13.8 DACcup ROSrub 525.5 100.0 0.0 23.1 20.9 463.8 72 436.1 91.7 42.3 PHYtri ACAcae 100.0 458.9 20.5 11.1 100.0 5.4 632.0 66.7 398.0 16.9 58.1 100.0 PRUfer 563.5 91.7 448.8 73 DIStou 15.0 8.6 100.0 16.4 31.9 87.5 75.0 506.9 PRUtax POMkum 64.4 1.0 349.3 36.5 100.0 92.9 487.8 13.4 3.0 230.0 100.0 100.0 19.3 AGAaus 74 CORarb 45.6 492.3 18.4 178.3 100.0 6.0 66.7 660.4 10.5 CUPmac 77.5 Asparagales 100.0 10.4 CORlae 42.8 193.4 75.0 92.3 462.8 12.7 PSEmen 75 53.3 433.0 11.1 18.6 21.4 Poaceae 100.0 319.1 8.8 QUEile 87.5 PINwal 3.8 455.0 87.5 388.2 13.8 18.2 75.0 19.7 76 BETpen 683.4 87.5 323.7 PINari 26.7 100.0 20.0 52.1 1.5 100.0 LOPmen 654.3 Poales 460.2 42.1 PINnig 45.8 23.9 100.0 69.2 29.4 77 235.5 100.0 Rosaceae 482.4 FUScli 90.9 86.0 31.3 PINsyl 59.9 16.3 460.3 Podocarpaceae 536.4 50.0 87.5 33.5 43.9 PINpin 78 FUSfus 46.9 60.6 490.0 100.0 85.7 699.7 Rosales 534.9 38.3 15.0 ACAdea 13.1 39.8 694.6 100.0 100.0 PINcon 539.8 28.0 5.0 79 8.2 619.9 100.0 93.3 PINpal TRIrep 86.7 3.6 299.9 95.2 100.0 495.0 23.8 41.7 PINrad 80 12.5 100.0 224.6 TRIarv 673.7 80.9 127.8 18.2 0.5 37.5 595.3 PINpon CARaus 0.0 187.0 100.0 Fagales 18.6 81 0.0 Pinales 100.0 63.1 64.2 12.5 371.6 188.0 PINcou SOPmic 150.0 79.5 22.5 93.2 0.0 Pinaceae 758.3 10.5 651.0 100.0 80.8 BLEpen 82 LUParb 2.3 100.0 552.0 25.0 4.4 225.6 75.0 0.4 POLves 40.0 213.8 11.0 83 ULEeur 11.3 25.0 3.8 83.8 210.9 Fabales PTEesc 50.0 330.2 28.6 CHApal 65.1 704.8 60.0 24.9 18.1 100.0 507.7 84 19.9 100.0 55.0 DICsqu CYTsco 314.8 100.0 606.3 21.5 31.3 100.0 CYAmed 13.3 342.3 19.1 28.8 85 MAYboa 87.5 62.5 388.1 26.7 7.3 28.1 CYAdea 399.7 100.0 87.5 442.3 10.0 ARIfru 86 7.9 2.3 200.9 62.5 0.0 150.0 0.0 0.0 LYCfas ARIser 8.6 2.1 248.2 42.1 IF (%) MT(°C) BT(s) BB(%) Code 87 100.0 Malpighiales 100.0 WEIrac 16.9 9.1 548.5 557.5 38.9 35.0 OLEpan 100.0 100.0 12.0 14.9 269.1 553.6 18.9 88 MELcra 100.0 27.5 OLEfur 339.5 65.2 394.8 17.6 6.4 11.3 17.1 MELram 0.0 20.0 204.7 OLEtra 89 0.0 150.0 0.6 0.0 100.0 0.0 19.0 VIOcun 347.1 150.0 CELgra 16.3 0.0 0.0 55.0 100.0 0.0 90 428.8 150.0 BRAlon SALfra 7.0 66.7 Asteraceae 0.0 0.0 23.3 150.0 0.0 334.2 0.0 0.0 LAGcun 91 SALmat 4.5 36.8 150.0 5.8 264.9 Sapindales 100.0 0.0 POPtri 3.3 100.0 598.5 0.0 LEPpec 446.5 0.0 92 14.7 100.0 33.0 0.0 5.7 12.5 150.0 FARjap POPnig 358.5 50.8 15.8 100.0 100.0 173.4 0.0 93 11.8 372.0 37.5 Asterales BRArep CHOter 18.3 0.0 491.2 0.5 0.0 12.7 221.5 100.0 HELfil NEMsqu 15.0 0.0 150.0 7.4 0.6 94 3.0 456.0 90.5 RAOgra 0.0 150.0 0.0 80.0 CITlim 4.4 11.0 100.0 408.6 0.0 150.0 RAOsub 95 36.4 Myrtales 25.0 0.0 0.0 DYSspe 20.0 9.3 551.9 150.0 69.2 100.0 0.0 0.0 HYPrad 29.5 21.9 234.0 206.3 AEShip 25.0 100.0 0.0 CREcap 96 2.2 285.6 250.3 0.0 63.8 100.0 0.0 DODvis 250.3 532.9 9.9 0.0 HIEpil 6.3 5.3 25.0 52.0 97 0.0 100.0 150.0 6.0 ALEexc 1.0 609.2 21.9 HIEpra 14.9 Apiales 100.0 239.1 14.0 222.5 12.5 11.3 PLAreg 15.8 HIElep 98 13.8 150.0 38.5 61 513.5 0.0 0.6 63 .5 66.8 HOHang 100.0 626.2 3.9 CORcot 69.4 188.3 85 .3 0.0 85.7 Ericales Lamiales 293.9 0.0 99 KELdie 233.9 88.2 .7 17.7 CORbud 23.1 0.4 100.0 100.0 270.6 613.0 Ericaceae 18.0 6.8 PIMore 0.0 100.0 62 384.6 WAHalb 4.4 397.6 Gentianales 59.2 50.0 .
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