Philippa Kaur

Delhey, K. A review of Gloger’s rule, an ecogeographical rule of colour: definitions, interpretations and evidence. Biol. Rev. 94, 1294–1316 (2019).PubMed 

Google Scholar 
Tian, L. & Benton, M. J. Predicting biotic responses to future climate warming with classic ecogeographic rules. Curr. Biol. 30, R744–R749 (2020).CAS 
PubMed 

Google Scholar 
Ryding, S., Klaassen, M., Tattersall, G. J., Gardner, J. L. & Symonds, M. R. E. Shape-shifting: changing animal morphologies as a response to climatic warming. Trends Ecol. Evol. 36, 1036–1048 (2021).Salewski, V. & Watt, C. Bergmann’s rule: a biophysiological rule examined in birds. Oikos 126, 161–172 (2017).
Google Scholar 
Allen, J. A. The influence of physical conditions in the genesis of species. Radic. Rev. 1, 108–140 (1877).
Google Scholar 
Ashton, K. G., Tracy, M. C. & De Queiroz, A. Is Bergmann’s rule valid for mammals? Am. Nat. 156, 390–415 (2000).PubMed 

Google Scholar 
Ashton, K. G. Patterns of within-species body size variation of birds: strong evidence for Bergmann’s rule. Glob. Ecol. Biogeogr. 11, 505–523 (2002).
Google Scholar 
Nudds, R. L. & Oswald, S. A. An interspecific test of Allen’s rule: evolutionary implications for endothermic species. Evolution (N. Y) 61, 2839–2848 (2007).CAS 

Google Scholar 
Symonds, M. R. E. & Tattersall, G. J. Geographical variation in bill size across bird species provides evidence for Allen’s rule. Am. Nat. 176, 188–197 (2010).PubMed 

Google Scholar 
Cardilini, A. P. A., Buchanan, K. L., Sherman, C. D. H., Cassey, P. & Symonds, M. R. E. Tests of ecogeographical relationships in a non-native species: what rules avian morphology? Oecologia 181, 783–793 (2016).ADS 
PubMed 

Google Scholar 
Alhajeri, B. H., Fourcade, Y., Upham, N. S. & Alhaddad, H. A global test of Allen’s rule in rodents. Glob. Ecol. Biogeogr. 29, 2248–2260 (2020).
Google Scholar 
McNab, B. K. On the ecological significance of Bergmann’s rule. Ecology 52, 845–854 (1971).
Google Scholar 
Meiri, S., Dayan, T. & Simberloff, D. Carnivores, biases and Bergmann’s rule. Biol. J. Linn. Soc. 81, 579–588 (2004).
Google Scholar 
Gohli, J. & Voje, K. L. An interspecific assessment of Bergmann’s rule in 22 mammalian families. BMC Evol. Biol. 16, 1–12 (2016).
Google Scholar 
Freeman, B. G. Little evidence for Bergmann’s rule body size clines in passerines along tropical elevational gradients. J. Biogeogr. 44, 502–510 (2017).
Google Scholar 
Riemer, K., Guralnick, R. P. & White, E. No general relationship between mass and temperature in endothermic species. Elife 7, e27166 (2018).PubMed 
PubMed Central 

Google Scholar 
Blackburn, T. M., Gaston, K. J. & Loder, N. Geographic gradients in body size: a clarification of Bergmann’s rule. Divers. Distrib. 5, 165–174 (1999).
Google Scholar 
Watt, C., Mitchell, S. & Salewski, V. Bergmann’s rule; a concept cluster? Oikos 119, 89–100 (2010).
Google Scholar 
James, F. C. Geographic size variation in birds and its relationship to climate. Ecology 51, 365–390 (1970).
Google Scholar 
Cartar, R. V. & Morrison, R. I. G. Metabolic correlates of leg length in breeding arctic shorebirds: the cost of getting high. J. Biogeogr. 32, 377–382 (2005).
Google Scholar 
Friedman, N. R., Harmáčková, L., Economo, E. P. & Remeš, V. Smaller beaks for colder winters: thermoregulation drives beak size evolution in Australasian songbirds. Evolution (N. Y). 71, 2120–2129 (2017).Fan, L., Cai, T., Xiong, Y., Song, G. & Lei, F. Bergmann’s rule and Allen’s rule in two passerine birds in China. Avian. Res. 10, 1–11 (2019).
Google Scholar 
Romano, A., Séchaud, R. & Roulin, A. Geographical variation in bill size provides evidence for Allen’s rule in a cosmopolitan raptor. Glob. Ecol. Biogeogr. 29, 65–75 (2020).
Google Scholar 
Romano, A., Séchaud, R. & Roulin, A. Generalized evidence for Bergmann’s rule: body size variation in a cosmopolitan owl genus. J. Biogeogr. 48, 51–63 (2021).
Google Scholar 
Gardner, J. L. et al. Spatial variation in avian bill size is associated with humidity in summer among Australian passerines. Clim. Chang. Responses 3, 1–11 (2016).
Google Scholar 
Greenberg, R. & Danner, R. M. The influence of the california marine layer on bill size in a generalist songbird. Evolution (N. Y) 66, 3825–3835 (2012).
Google Scholar 
Greenberg, R., Danner, R., Olsen, B. & Luther, D. High summer temperature explains bill size variation in salt marsh sparrows. Ecography (Cop.) 35, 146–152 (2012).
Google Scholar 
Klir, J. J. & Heath, J. E. An infrared thermographic study of surface temperature in relation to external thermal stress in three species of foxes: the red fox (Vulpes vulpes), Arctic fox, and kit fox (Vulpes macrotis). Physiol. Zool. 65, 1011–1021 (1992).
Google Scholar 
Ballentine, B. & Greenberg, R. Common garden experiment reveals genetic control of phenotypic divergence between swamp sparrow subspecies that lack divergence in neutral genotypes. PLoS One 5, 1–6 (2010).
Google Scholar 
Nord, A. & Giroud, S. Lifelong effects of thermal challenges during development in birds and mammals. Front. Physiol. 11, 1–9 (2020).
Google Scholar 
Riek, A. & Geiser, F. Developmental phenotypic plasticity in a marsupial. J. Exp. Biol. 215, 1552–1558 (2012).PubMed 

Google Scholar 
Cunningham, S. J., Martin, R. O., Hojem, C. L. & Hockey, P. A. R. Temperatures in excess of critical thresholds threaten nestling growth and survival in a rapidly-warming arid savanna: a study of common fiscals. PLoS One 8, e74613 (2013).Mariette, M. M. & Buchanan, K. L. Prenatal acoustic communication programs offspring for high posthatching temperatures in a songbird. Science 353, 812–814 (2016).ADS 
CAS 
PubMed 

Google Scholar 
Nord, A. & Nilsson, J. Å. Incubation temperature affects growth and energy metabolism in blue tit nestlings. Am. Nat. 178, 639–651 (2011).PubMed 

Google Scholar 
Serrat, M. A. Allen’s rule revisited: temperature influences bone elongation during a critical period of postnatal development. Anat. Rec. 296, 1534–1545 (2013).
Google Scholar 
Larson, E. R. et al. Nest microclimate predicts bill growth in the Adelaide rosella (Aves: Psittaculidae). Biol. J. Linn. Soc. 124, 339–349 (2018).
Google Scholar 
Burness, G., Huard, J. R., Malcolm, E. & Tattersall, G. J. Post-hatch heat warms adult beaks: irreversible physiological plasticity in Japanese quail. Proc. R. Soc. B Biol. Sci. 280, 20131436 (2013).Husby, A., Hille, S. M. & Visser, M. E. Testing mechanisms of bergmann’s rule: phenotypic decline but no genetic change in body size in three passerine bird populations. Am. Nat. 178, 202–213 (2011).PubMed 

Google Scholar 
Cresswell, W., Clark, J. A. & Macleod, R. How climate change might influence the starvation-predation risk trade-off response. Proc. R. Soc. B Biol. Sci. 276, 3553–3560 (2009).CAS 

Google Scholar 
McNamara, J. M., Higginson, A. D. & Verhulst, S. The influence of the starvation-predation trade-off on the relationship between ambient temperature and body size among endotherms. J. Biogeogr. 43, 809–819 (2016).PubMed 

Google Scholar 
Dickman, C. R. Body size, prey size, and community structure in insectivorous mammals. Ecology 69, 569–580 (1988).
Google Scholar 
Carbone, C., Mace, G. M., Roberts, S. C. & Macdonald, D. W. Energetic constraints on the diet of terrestrial carnivores. Nature 402, 286–288 (1999).ADS 
CAS 
PubMed 

Google Scholar 
Cohen, J. E., Pimm, S. L., Yodzis, P., & Saldaña, J. Body sizes of animal predators and animal prey in food webs. J. Anim. Ecol. 62, 67–78 (1993).
Google Scholar 
McKinnon, L. et al. Lower predation risk for migratory birds at high latitudes. Science 327, 326–327 (2010).ADS 
CAS 
PubMed 

Google Scholar 
Díaz, M. et al. The geography of fear: a latitudinal gradient in anti-predator escape distances of birds across Europe. PLoS One 8, e64634 (2013).Gosler, A. G., Greenwood, J. J. D. & Perrins, C. Predation risk and the cost of being fat. Nature 377, 621–623 (1995).ADS 
CAS 

Google Scholar 
Anderson, A. M. et al. Consistent declines in wing lengths of Calidridine sandpipers suggest a rapid morphometric response to environmental change. PLoS One 14, 1–21 (2019).CAS 

Google Scholar 
Milá, B., Wayne, R. K. & Smith, T. B. Ecomorphology of migratory and sedentary populations of the yellow-rumped warbler (Dendroica Coronata). Condor 110, 335–344 (2008).
Google Scholar 
O’Hara, P. D., Fernández, G., Haase, B., de la Cueva, H. & Lank, D. B. Differential migration in western sandpipers with respect to body size and wing length. Condor 108, 225–232 (2006).
Google Scholar 
Ketterson, E. D. & Nolan, V. Geographic variation and its climatic correlates in the sex ratio of eastern-wintering dark-eyed juncos (Junco hyemalis hyemalis). Ecology 57, 679–693 (1976).
Google Scholar 
Nebel, S. Differential migration of shorebirds in the East Asian-Australasian Flyway. Emu 107, 14–18 (2007).
Google Scholar 
Elner, R. W. & Seaman, D. A. Calidrid conservation: unrequited needs. Wader Study Gr. Bull. 100, 30–34 (2003).
Google Scholar 
Greenberg, R. Dissimilar bill shapes in new world tropical versus temperate forest foliage-gleaning birds. Oecologia 49, 143–147 (1981).ADS 
PubMed 

Google Scholar 
Nebel, S. Latitudinal clines in bill length and sex ratio in a migratory shorebird: a case of resource partitioning? Acta Oecologica 28, 33–38 (2005).ADS 

Google Scholar 
Mathot, K. J., Smith, B. D. & Elner, R. W. Latitudinal clines in food distribution correlate with differential migration in the Western Sandpiper. Ecology 88, 781–791 (2007).PubMed 

Google Scholar 
Duijns, S. et al. Sex-specific winter distribution in a sexually dimorphic shorebird is explained by resource partitioning. Ecol. Evol. 4, 4009–4018 (2014).PubMed 
PubMed Central 

Google Scholar 
Wilson, J. R., Nebel, S. & Minton, C. D. T. Migration ecology and morphometrics of two Bar-tailed Godwit populations in Australia. Emu 107, 262–274 (2007).
Google Scholar 
Nebel, S., Rogers, K. G., Minton, C. D. T. & Rogers, D. I. Is geographical variation in the size of Australian shorebirds consistent with hypotheses on differential migration? Emu 113, 99–111 (2013).
Google Scholar 
Beltran, R. S., Burns, J. M. & Breed, G. A. Convergence of biannual moulting strategies across birds and mammals. Proc. R. Soc. B Biol. Sci. 285, 20180318 (2018).Tattersall, G. J., Arnaout, B. & Symonds, M. R. E. The evolution of the avian bill as a thermoregulatory organ. Biol. Rev. 92, 1630–1656 (2017).PubMed 

Google Scholar 
Battley, P. F., Rogers, D. I., Piersma, T. & Koolhaas, A. Behavioural evidence for heat-load problems in Great Knots in tropical Australia fuelling for long-distance flight. Emu 103, 97–103 (2003).
Google Scholar 
Rogers, D. I., Piersma, T. & Hassell, C. J. Roost availability may constrain shorebird distribution: Exploring the energetic costs of roosting and disturbance around a tropical bay. Biol. Conserv. 133, 225–235 (2006).
Google Scholar 
Danner, R. M. & Greenberg, R. A critical season approach to Allen’s rule: Bill size declines with winter temperature in a cold temperate environment. J. Biogeogr. 42, 114–120 (2015).
Google Scholar 
Buchholz, R. Thermoregulatory role of the unfeathered head and neck in male wild turkeys. Auk 113, 310–318 (1996).
Google Scholar 
Marchant, S. & Higgins, P. J. (eds.) Handbook of Australian, New Zealand and Antarctic Birds. Volume 2: Raptors to Lapwings (Oxford University Press, 1993).Higgins, P. J. & Davies, S. J. J. F. (eds.) Handbook of Australian, New Zealand and Antarctic Birds. Volume 3: Snipe to Pigeons (Oxford University Press, 1996).Andrew, S. C., Hurley, L. L., Mariette, M. M. & Griffith, S. C. Higher temperatures during development reduce body size in the zebra finch in the laboratory and in the wild. J. Evol. Biol. 30, 2156–2164 (2017).CAS 
PubMed 

Google Scholar 
Morrick, Z. N. et al. Differential population trends align with migratory connectivity in an endangered shorebird. Conserv. Sci. Pract. 4, 1–13 (2022).
Google Scholar 
Hassell, C., Southey, I., Boyle, A. & Yang, H.-Y. Red knot Calidris canutus: subspecies and migration in the East Asian-Australasian flyway – where do all the red knot go? BirdingASIA 16, 89–93 (2011).
Google Scholar 
Battley, P. F. et al. Contrasting extreme long-distance migration patterns in bar-tailed godwits Limosa lapponica. J. Avian Biol. 43, 21–32 (2012).
Google Scholar 
Aharon-Rotman, Y., Buchanan, K. L., Clark, N. J., Klaassen, M. & Buttemer, W. A. Why fly the extra mile? Using stress biomarkers to assess wintering habitat quality in migratory shorebirds. Oecologia 182, 385–395 (2016).ADS 
PubMed 

Google Scholar 
Aharon-Rotman, Y., Gosbell, K., Minton, C. & Klaassen, M. Why fly the extra mile? Latitudinal trend in migratory fuel deposition rate as driver of trans-equatorial long-distance migration. Ecol. Evol. 6, 6616–6624 (2016).PubMed 
PubMed Central 

Google Scholar 
Hollands, D. & Minton, C. Waders: The Shorebirds of Australia (Bloomings Books, 2012).Siepielski, A. M. et al. No evidence that warmer temperatures are associated with selection for smaller body sizes. Proc. R. Soc. B Biol. Sci. 286, 20191332 (2019).Ho, C. K., Pennings, S. C. & Carefoot, T. H. Is diet quality an overlooked mechanism for Bergmann’s rule? Am. Nat. 175, 269–276 (2010).PubMed 

Google Scholar 
Piersma, T. et al. Fuel storage rates in Red Knots worldwide: facing the severest ecological constraint in tropical intertidal environments? In Birds of Two Worlds: Ecology and Evolution of Migration (eds Greenburg, R. & Marra, P. P.) (Smithsonian Institution Press, 2005).Hedenström, A. & Rosén, M. Predator versus prey: on aerial hunting and escape strategies in birds. Behav. Ecol. 12, 150–156 (2001).
Google Scholar 
Van Den Hout, P. J., Mathot, K. J., Maas, L. R. M. & Piersma, T. Predator escape tactics in birds: linking ecology and aerodynamics. Behav. Ecol. 21, 16–25 (2010).
Google Scholar 
Schemske, D. W., Mittelbach, G. G., Cornell, H. V., Sobel, J. M. & Roy, K. Is there a latitudinal gradient in the importance of biotic interactions? Annu. Rev. Ecol. Evol. Syst. 40, 245–269 (2009).
Google Scholar 
Cain, K. E. et al. Conspicuous plumage does not increase predation risk: a continent-wide test using model songbirds. Am. Nat. 193, 359–372 (2019).PubMed 

Google Scholar 
Cohen, J. E., Pimm, S. L., Yodzis, P. & Saldana, J. Body sizes of animal predators and animal prey in food webs. J. Anim. Ecol. 62, 67–78 (1993).
Google Scholar 
Gotmark, F. & Post, P. Prey selection by sparrowhawks, Accipiter nisus: relative predation risk for breeding passerine birds in relation to their size, ecology and behaviour. Philos. Trans. R. Soc. B Biol. Sci. 351, 1559–1577 (1996).ADS 

Google Scholar 
McQueen, A. et al. Evolutionary drivers of seasonal plumage colours: colour change by moult correlates with sexual selection, predation risk and seasonality across passerines. Ecol. Lett. 22, 1838–1849 (2019).PubMed 

Google Scholar 
Martínez, A. E. & Zenil, R. T. Foraging guild influences dependence on heterospecific alarm calls in Amazonian bird flocks. Behav. Ecol. 23, 544–550 (2012).
Google Scholar 
Gauthreaux, S. A. The ecological significance of behavioral dominance. In Social Behavior. Perspectives in Ethology, vol 3 (eds Bateson, P. P. G. & Klopfer, P. H.) (Springer, 1978).Friedman, N. R. et al. Evolution of a multifunctional trait: Shared effects of foraging ecology and thermoregulation on beak morphology, with consequences for song evolution. Proc. R. Soc. B Biol. Sci. 286, 20192474 (2019).Campbell-Tennant, D. J. E., Gardner, J. L., Kearney, M. R. & Symonds, M. R. E. Climate-related spatial and temporal variation in bill morphology over the past century in Australian parrots. J. Biogeogr. 42, 1163–1175 (2015).
Google Scholar 
Sullivan, T. N., Meyers, M. A. & Arzt, E. Scaling of bird wings and feathers for efficient flight. Sci. Adv. 5, 1–9 (2019).
Google Scholar 
Gosler, A. G., Greenwood, J. J. D., Baker, J. K. & Davidson, N. C. The field determination of body size and condition in passerines: a report to the British Ringing Committee. Bird. Study 45, 92–103 (1998).
Google Scholar 
Tattersall, G. J., Chaves, J. A. & Danner, R. M. Thermoregulatory windows in Darwin’s finches. Funct. Ecol. 32, 358–368 (2018).
Google Scholar 
Weeks, B. C. et al. Shared morphological consequences of global warming in North American migratory birds. Ecol. Lett. 23, 316–325 (2020).PubMed 

Google Scholar 
Minton, C. The history and achievements of the Victorian Wader Study Group. Stilt 50, 285–294 (2006).
Google Scholar 
Minton, C. The history of wader studies in north-west Australia. Stilt 50, 224–234 (2006).
Google Scholar 
Lowe, K. W. The Australian Bird Bander’s Manual (Australian Bird and Bat Banding Scemes, Australian National Parks and Wildlife Services, 1989).Aarif, K. M. Some aspects of feeding ecology of the lesser sand plover Charadrius mongolus in three different zones in the Kadalundy Estuary, Kerala, South India. Podoces 4, 100–1007 (2009).
Google Scholar 
Bates, D., Maechler, M. & Bolker, B. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).
Google Scholar 
Rue, H. et al. Bayesian computing with INLA: a review. Annu. Rev. Stat. Its Appl. 4, 395–421 (2017).ADS 

Google Scholar 
Li, D., Dinnage, R., Nell, L. A., Helmus, M. R. & Ives, A. R. phyr: an r package for phylogenetic species-distribution modelling in ecological communities. Methods Ecol. Evol. 11, 1455–1463 (2020).
Google Scholar 
Simpson, D., Rue, H., Riebler, A., Martins, T. G. & Sørbye, S. H. Penalising model component complexity: a principled, practical approach to constructing priors. Stat. Sci. 32, 1–28 (2017).MathSciNet 
MATH 

Google Scholar 
Jetz, W., Thomas, G. H., Joy, J. B., Hartmann, K. & Mooers, A. O. The global diversity of birds in space and time. Nature 491, 444–448 (2012).ADS 
CAS 
PubMed 

Google Scholar 
Schliep, K. Phangorn: phylogenetic analysis in R. Bioinformatics 27, 592–593 (2011).CAS 
PubMed 

Google Scholar 
McQueen, A et al. Data from: thermal adaptation best explains Bergmann’s and Allen’s rule across ecologically diverse shorebirds. Dryad Dataset. https://doi.org/10.5061/dryad.xsj3tx9j5.Tattersall, G. J., Andrade, D. V. & Abe, A. S. Heat exchange from the toucan bill reveals a controllable vascular thermal radiator. Science 325, 468–470 (2009).ADS 
CAS 
PubMed 

Google Scholar 
Greenberg, R., Cadena, V., Danner, R. M. & Tattersall, G. Heat loss may explain bill size differences between birds occupying different habitats. PLoS One 7, 1–9 (2012).
Google Scholar 
Ryeland, J., Weston, M. A. & Symonds, M. R. E. Bill size mediates behavioural thermoregulation in birds. Funct. Ecol. 31, 885–893 (2017).
Google Scholar 
Pavlovic, G., Weston, M. A. & Symonds, M. R. E. Morphology and geography predict the use of heat conservation behaviours across birds. Funct. Ecol. 33, 286–296 (2019).
Google Scholar  More

Dinosauria—Owen, 184225,Ornithischia—Seeley, 188726,Thyreophora—Nopcsa, 191527,Jakapil kaniukura gen. et sp. nov. (Figs. 1, 2, 3, 4, Suppl. Figs. 2, 3).Figure 1Holotype of Jakapil kaniukura (MPCA-PV-630), skull bones. (a) Skull bones in right lateral view (dashed contours based on Scelidosaurus10); (b) basisphenoid in left lateral view. af anterior foramen, btp basipterygoid process, bt basal tubera, cp cultriform process, df double foramen, ene external naris edge, jf jugal facet of the maxilla, Mx maxilla, mxe maxillary emargination, Pmx premaxilla, vc Vidian canal, vp ventral process.Full size imageFigure 2Holotype of Jakapil kaniukura (MPCA-PV-630), lower jaw bones. (a) left mandible in lateral view; (b) left mandible in lateral view, interpreted bone contours; (c) left mandible in medial view; (d) left mandible in medial view, interpreted bone contours; (e) right surangular in lateral view (mirrored); (f) transversal section of the posterior half of the left mandible, cranial view; (g) articular bone in occlusal view; (h) predentary bone in occlusal view. A angular, af adductor fossa, Ar articular, Ar (gl) glenoid fossa of the articular, ce coronoid eminence, D dentary, de dentary emargination, dfo dentary foramen, dmp dorsomedial process of the articular, dr dentary rugosities, hi subhorizontal inflection (dashed line), imf internal mandibular fenestra, lp lateral process of the predentary, mc Meckelian canal, Pa prearticular, Pd predentary, rp retroarticular process, S surangular, saf surangular facet for the glenoid articulation, safo surangular foramen (canal), Sp splenial, st surangular tubercle, sy mandibular symphysis, vmc ventral mandibular crest.Full size imageFigure 3Holotype of Jakapil kaniukura (MPCA-PV-630), teeth. Maxillary teeth in labial (a,b) and lingual (c,d); (d) highlight the wear facet) views; dentary teeth in lingual (e,g–j); (h,j) highlight the wear facets) and labial (f) views. dwf dentary tooth wear facet, me prominent mesial edge, mwf maxillary tooth wear facet.Full size imageFigure 4Holotype of Jakapil kaniukura (MPCA-PV-630), postcranial bones. Speculative silhouette showing preserved elements (a); osteoderm distribution is speculative and partial to show non-osteodermal elements); dorsal vertebra elements in dorsal (b), right lateral (c) and anterior (d,e) views; sacral vertebra in left lateral view (f); mid-caudal vertebra in left lateral view (g); fragment of the mid-shaft of a dorsal rib in posterior view (the enlarged, broken posterior edge is highlighted (h); expanded distal ends of two dorsal ribs (i); left scapula in lateral view (j); right scapula in lateral view (k); right coracoid in lateral view (l); left and right humeri in anterior view (m); probable right ulna in lateral view (n); metacarpals, non-ungual and ungual phalanx in dorsal views (o); left femur elements in anterior view (p); proximal end of the right fibula in lateral view (q); distal end of the left tibia in anterior view (r); ischial elements in side view (s); cervical osteoderms in dorsal view (t), flat scutes in dorsal view (u), spine-like osteoderm in side view (v) and ossicle in dorsal view (w). ac acromial crest, aco asymmetrical cervical osteoderm, alp anterolateral process, ap acromial process, at anterior trochanter, bb basal bone, ebr expanded broken rib edge, di diapophysis, dpc deltopectoral crest, ft fourth trochanter, gl glenoid, mc metacarpals, nc neural canal, ncs neurocentral suture, ph non-ungual phalanx, pp pubic peduncle, poz postzygapophyses, rug marginal rugosities, sb scapular blade, sc scute, tp transverse process, uph ungual phalanx.Full size imageEtymologyThe genus, Jakapil (Ja-Kapïl: shield bearer), comes from the ‘gananah iahish’, Puelchean or northern Tehuelchean language. The specific epithet, comprising kaniu (crest) and kura (stone), refers to the diagnostic ventral crest of the mandible, and comes from the Mapudungun language. These languages, currently spoken by more than 200,000 people, have been combined as a tribute to both of the coexisting native populations of North Patagonia, South America.HolotypeMPCA-PV-630 is a partial skeleton of a subadult individual (see Supplementary Information) that preserves fragments of some cranial bones (premaxilla, maxilla and basisphenoid), approximately 15 partial teeth and fragments, a nearly complete left lower jaw plus an isolated surangular, 12 partial vertebral elements, a complete dorsal rib and fifteen rib fragments, a partial coracoid, a nearly complete left scapula, a partial right scapula, two partial humeri, a possible partial right ulna, a complete and a partial metacarpal bone, three ischial and two femoral fragments, the distal end of a right tibia, the proximal end of a right fibula, three pedal phalanges, and more than forty osteoderms.Referred specimensMPCA-PV-371, two partial conical osteoderms.Locality and horizonUpper beds of the Candeleros Formation, early Late Cretaceous (Cenomanian, ~ 94–97 My, see16, and references therein), locality of Cerro Policía, Río Negro Province, North Patagonia, Argentina (Suppl. Fig. 1).DiagnosisJakapil differs from all other thyreophorans in having: a large, ventral crest on the posterior half of the lower jaw, which is composed of the dentary, the angular and the splenial (medially hidden by the crest); a dorsomedially directed process in the short retroarticular process; leaf-shaped tooth crowns with a prominent mesial edge on their labial surface; maxillary and dentary tooth crowns differ from each other in their apical contour, the former being pointed and strongly asymmetrical, and the latter slightly curved distally with a more rounded and less asymmetrical contour; elongated (articular surface almost or completely beyond the posterior centrum face) and slender (width of less than a half postzygapophyses length) postzygapophyses in dorsal vertebrae; a strongly reduced humerus relative to the femur (proximal humeral width smaller than distal femoral width, see Supplementary Information), with a deep proximal fossa distally delimited by a curved ridge; a very large fibula relative to the femur (anteroposterior length of the proximal end almost comparable to the distal width of the femur); flattened and thin disk-like postcranial osteoderms.Summarized descriptionA detailed description of the holotype is provided in the Supplementary Information. Jakapil is a small thyreophoran dinosaur (the subadult holotype is estimated to have been less than 1.5 m in body length and to have weighed 4.5–7 kg; see Supplementary Information, femoral description), with several novelties for a thyreophoran dinosaur.A short skull is suggested by the size of the skull and jaw bones, and the reduced number of dentary tooth positions (eleven), compared with most non-ankylosaurid thyreophorans28,29. The antorbital and mandibular fenestrae seem absent, as in ankylosaurs29 (Fig. 1a; the mandibular fenestra is also absent in Scelidosaurus10). Dentary and maxillary emarginations are present, as usual in ornithischians30 (Fig. 1a). The block-like basisphenoid is strongly similar to that of Scelidosaurus10, with Vidian canals opened posterodorsally to the basipterygoid processes, the basipterygoid processes lateroventrally projected (unlike the anteriorly directed processes of stegosaurs28 and ankylosaurs29), and a strong cultriform process (as in Lesothosaurus31, Thescelosaurus32 and probably Scelidosaurus10; Fig. 1b).Jakapil also bears the first predentary bone (Fig. 2a–d) with a plesiomorphic shape in a thyreophoran. It is subtriangular and quite similar to that of Lesothosaurus31, and externally it is ornamented by sulci and foramina, suggesting the presence of a keratinous beak. A beak is also supported in the edentulous and subtly ornamented preserved part of the premaxilla, as in derived thyreophorans28,29. The posterior half of the short lower jaw (Fig. 2a–f) is strongly dorsoventrally expanded, resembling the general shape of the heterodontosaurid33 and basal ceratopsian jaws34. This expansion is composed of a well-developed coronoid eminence (Fig. 2a–d, ce; similar to that in the stegosaur Huayangosaurus35 and most ankylosaurs36) and a large ventral crest at the dentary-angular contact that is unique among thyreophorans (Fig. 2a–d,f, vmc; resembling that of some ceratopsians, see SI). The dentary symphysis is slightly spout-shaped, as in most ornithischians37. Anteriorly, the dentary oral margin is subhorizontal in lateral view (Fig. 2a–d, D), unlike the strongly downturned line of most thyreophorans30,37. There is no evidence of a mandibular osteoderm as occurs in Scelidosaurus and ankylosaurs10. A surangular tubercle (Fig. 2a, st) adjacent to the glenoid fossa seems anteriorly continued by a subtly developed subhorizontal inflection of the anterior lamina (Fig. 2e, hi), in the position of the surangular ridge (synapomorphy of Thyreophora37), though the first is poorly developed. The glenoid fossa is roughly aligned with the tooth row in lateral view (Fig. 2a–d). The short retroarticular process bears a dorsomedially directed process resembling that of several theropods (Fig. 2g, dmp; see Discussion). This process is absent in all other thyreophorans 9,10,35,36.The tooth crowns are leaf-shaped as in basal ornithischian and thyreophorans10,28,29,38 (Fig. 3). The tooth crowns are swollen labially at their base and lack both cingulum and ornamentation, unlike those of derived eurypodans28,29, heterodontosaurids33 and most neornithischians30,32. The mesial edge of the labial surface in the maxillary and dentary tooth crowns is prominent as in Scelidosaurus10, and ends distally in a denticle-like structure in Jakapil (Fig. 3, me). This prominent edge delimits anteriorly the wear facets of the dentary teeth. A striking difference with respect to most thyreophorans is that the maxillary and dentary tooth crowns are quite different (see Supplementary Information). The maxillary teeth (Fig. 3a–d) show seven/eight mesial and four distal denticles, a vertical apical denticle, and a straighter mesial denticle row (resembling those of non-ankylosaurid and non-stegosaurid thyreophorans10,35,36). The dentary teeth (Fig. 3e–j) bear seven mesial and five/six distal denticles, and a distally curved apical-most denticle. Also, the mesial denticle row is lingually recurved, as in Huayangosaurus35. Large, high-angled wear facets are present (Fig. 3d,h,j; dwf and mwf).The axial elements are similar to those of Scelidosaurus39 (Fig. 4). The posterior articular surface of an isolated cervical centrum is flattened and seems almost as wide as high. A large foramen is placed just posteroventral to the parapophysis. The dorsal centra are cylindrical and elongated, with subcircular articular surfaces, and are biconcave (Fig. 4c,e). The neural arch is low but the neural canal is larger (Fig. 4d,e, nc). A dorsal neurocentral suture is visible (Fig. 4c, ncs). The diapophyses are laterodorsally directed almost 40° from the horizontal (Fig. 4d, di), at a lower angle than in stegosaurs28 and most ankylosaurs29, unlike the horizontal processes of basal ornithischians38. The postzygapophyses are medially fused in a slender (width of less than a half postzygapophyses length) and strongly elongated posteriorly structure (Fig. 4b, poz; more than in some ankylosaurs, such as Euoplocephalus and Polacanthus; see40,41). An isolated mid-caudal vertebra shows an equidimensional centrum in lateral view, with concave, oval articular surfaces (Fig. 4g). Transverse processes are very small and button-like (Fig. 4g, tp). Postzygapophyses are medially fused and do not extend beyond the centrum edge (Fig. 4g, poz). Proximally, the cross-section of the dorsal ribs is T-shaped. The low curvature of the shaft suggests a wide torso, as occurs in Emausaurus42, Scelidosaurus39, and ankylosaurs29. Some rib fragments with expanded (though broken) posterior edges suggest the presence of intercostal bones (Fig. 4h, ebr), as in Scelidosaurus39, Huayangosaurus43,44, some ankylosaurids45 (and references therein) and some basal ornithopods46. Some ribs are distally expanded (Fig. 4i) like the anterior dorsal ribs of Scelidosaurus39 and Huayangosaurus43.Girdle and limb bones (see also Suppl. Figs. 2, 3) are mostly broken and with boreholes (probably due to bioerosion) at their ends. The scapular blade (Fig. 4j, sb) is elongated and parallel-sided, without distal expansion, an overall shape that resembles that of several theropods47, contrasting the distally expanded condition in most ornithischians30. A straight and parallel sided scapular blade is common in ankylosaurids29,40. The proximal scapular plate with a high acromial process (Fig. 4j,k, ap) is stegosaurian-like, and the lateral acromial crest (Fig. 4j,k, ac) is developed as in Huayangosaurus43. A low distinct ridge rises posterior to the glenoid fossa and represents the insertion site for the muscle triceps longus caudalis, as occur in ankylosaurids 40. The incomplete coracoid (Fig. 4l) is much shorter than the scapula, unlike that of ankylosaurs29,40, which bear a large coracoid. The coracoid and the scapula are not fused. The partial humeri (Fig. 3m) are strongly reduced in size, with overall limb proportions resembling those of basal ornithischians3,38 and several theropods47. A possible proximal end of the ulna (Fig. 4n) resembles that of other basal ornithischians, though more strongly laterally compressed. The anterolateral process is present (Fig. 4n, alp), and the olecranon process seems absent or poorly developed, as in Scutellosaurus9 and Scelidosaurus39. The ischia are poorly preserved (Fig. 4s). The pubic peduncle is separated from the iliac articulation, unlike the continuous cup-shaped structure of most ankylosaurs29. The shaft of the ischium is straight and parallel-edged, as in Scutellosaurus9 and Scelidosaurus39, and distally tapers as in stegosaurs28. The preserved femoral pieces (Fig. 4p) resemble those of basal ornithischians38,39. The bases of both the broken anterior and fourth trochanters (Fig. 4p, at, ft) are large, suggesting large elements; the fourth trochanter is proximally placed on the femoral shaft (near the height of the base of the anterior trochanter); and the distal end of the femur is slightly curved posteriorly. The proximal end of the right fibula (Fig. 4q) is much larger than that of all other thyreophorans (compared with both the femoral and tibial distal ends) and bears a large anterior curved crest. The block-like non-ungual phalanges and a bluntly pointed hoof-like ungual (Fig. 4o, ph, uph) are similar to those of Scelidosaurus39.At least five osteoderm types are preserved in the holotype of Jakapil. The cervical elements are composed of an external, low-crested scute (Fig. 4t, sc) over a fused, smooth bone base (Fig. 4t, bb), as in Scelidosaurus48 and several ankylosaurs2,49. A probable cervical element is also composed of a concave base of smooth bone fused to a high, asymmetrical osteoderm (Fig. 4t, aco). The bases of these dermal elements present strong rugosities at one edge, suggesting a sutural contact between (Fig. 4t, rug), as in Scelidosaurus48 and some ankylosaurs (such as Pinacosaurus and Scolosaurus40,49,50). Scute-like post-cervical osteoderms (Fig. 4u) are strongly flattened, disk-shaped, and suboval with a very low crest, resembling those of few ankylosaurs such as Gastonia and Gargoyleosaurus51 (‘body osteoderms’ sensu Kinneer et al.52; see also49). Only one scute shows a high triangular cross-section like those of Scelidosaurus48. Also present are a few conical, spike-like osteoderms with deep concave bases (Fig. 4v), and many flat, disk-shaped, minute (7–10 mm) ossicles without crests (Fig. 4w).PhylogenyThe phylogenetic analysis using the matrix of Soto-Acuña et al.5 recovers Jakapil within Thyreophora, as the sister taxon of Ankylosauria (Fig. 5). The branch support for the basal thyreophorans is considerably lower than that obtained by Soto-Acuña et al.5, although the support of Stegosauria and some less inclusive eurypodan clades is slightly better (ceratopsians and pachycephalosaurs also show a lower support). The Jakapil autapomorphies in this analysis are: ventrally orientated basipterygoid processes (char. 134; shared with Agilisaurus, Hypsilophodon, Zalmoxes, Tenontosaurus, Dryosaurus, Liaoceratops, Yamaceratops, Leptoceratops, Bagaceratops and Protoceratops); lateral orientation of the basipterygoid process articular facet (char. 136; shared with Homalocephale, Prenocephale, Stegoceras and Yinlong); a straight dentary tooth row in lateral view (char. 166; shared with the ornithischians Lesothosaurus, Eocursor, Scutellosaurus, Pinacosaurus, Euoplocephalus, heterodontosaurids and neornithischians); the presence of a ventral flange on the dentary (char. 170; shared with Psittacosaurus, Yamaceratops and Protoceratops); a well-developed coronoid process (char. 174; shared with heterodontosaurids and neornithischians); a surangular length of more than 50% the mandibular length (char. 183; shared with Stegoceras, Psittacosaurus, Yinlong, Chaoyangsaurus and Hualianceratops); less than 15 dentary teeth (char. 204; shared with heterodontosaurids, Gasparinisaura, Hypsilophodon, Wannanosaurus, Tenontosaurus, Dryosaurus and ceratopsians); apicobasally tall and blade-like cheek teeth crowns (char. 205; shared with Laquintasaura, Psittacosaurus, Yinlong, Chaoyangsaurus and Hualianceratops). Alternative phylogenetic analyses using the data matrices of Maidment et al.4, Norman6 and Wiersma and Irmis8 recover Jakapil as the sister taxon of Eurypoda (Stegosauria + Ankylosauria) and as a basal ankylosaur, respectively (see Supplementary Information). Being recovered either as an ankylosauromorph or a stem-eurypodan, Jakapil is closely related to Scelidosaurus in all analyses. Detailed phylogenetic results and discussion are provided in the Supplementary Information.Figure 5Time-calibrated strict consensus of 26,784 most parsimonious trees (L = 1267) with the Soto-Acuña et al.5 matrix. CI 0.359, RI: 0.708. Branch supports are figured (Bremer/bootstrap). Record ages references are listed in the Supplementary Information (Suppl. Fig. 4).Full size image More

Richardson, D. M. et al. Naturalization and invasion of alien plants: concepts and definitions. Divers. Distrib. 6, 93–107 (2000).Article 

Google Scholar 
van Kleunen, M. et al. Global exchange and accumulation of non-native plants. Nature 525, 100–103 (2015).PubMed 
Article 
CAS 

Google Scholar 
Capinha, C., Essl, F., Seebens, H., Moser, D. & Pereira, H. M. The dispersal of alien species redefines biogeography in the Anthropocene. Science 348, 1248–1251 (2015).CAS 
PubMed 
Article 

Google Scholar 
Vilà, M. & Hulme, P. E. in Impact of Biological Invasions on Ecosystem Services Vol. 12 Invading Nature – Springer Series in Invasion Ecology (eds Vilà, M. & Hulme, P. E.) 1–14 (Springer, 2017).Pyšek, P. et al. A global assessment of invasive plant impacts on resident species, communities and ecosystems: the interaction of impact measures, invading species’ traits and environment. Glob. Chang. Biol. 18, 1725–1737 (2012).PubMed Central 
Article 

Google Scholar 
Pyšek, P. et al. Scientists’ warning on invasive alien species. Biol. Rev. 95, 1511–1534 (2020).PubMed 
Article 

Google Scholar 
Bacher, S. et al. Socio-economic impact classification of alien taxa (SEICAT). Methods Ecol. Evol. 9, 159–168 (2018).Article 

Google Scholar 
Seebens, H. et al. No saturation in the accumulation of alien species worldwide. Nat. Commun. 8, 14435 (2017).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Seebens, H. et al. Projecting the continental accumulation of alien species through to 2050. Glob. Chang. Biol. 27, 970–982 (2021).CAS 
Article 

Google Scholar 
Kriticos, D. J., Sutherst, R. W., Brown, J. R., Adkins, S. W. & Maywald, G. F. Climate change and the potential distribution of an invasive alien plant: Acacia nilotica ssp. indica in Australia. J. Appl. Ecol. 40, 111–124 (2003).Article 

Google Scholar 
Thuiller, W., Richardson, D. M. & Midgley, G. F. in Biological Invasions (ed. Nentwig, W.) 197–211 (Springer, 2007).Hobbs, R. J. in Invasive Species in a Changing World (eds Mooney, H. A. & Hobbs, R. J.) 55–64 (Island Press, 2000).Seebens, H. et al. Global trade will accelerate plant invasions in emerging economies under climate change. Glob. Chang. Biol. 21, 4128–4140 (2015).PubMed 
Article 

Google Scholar 
Razanajatovo, M. et al. Plants capable of selfing are more likely to become naturalized. Nat. Commun. 7, 13313 (2016).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Bucharova, A. & van Kleunen, M. Introduction history and species characteristics partly explain naturalization success of North American woody species in Europe. J. Ecol. 97, 230–238 (2009).Article 

Google Scholar 
Ordonez, A., Wright, I. J. & Olff, H. Functional differences between native and alien species: a global-scale comparison. Funct. Ecol. 24, 1353–1361 (2010).Article 

Google Scholar 
van Kleunen, M., Weber, E. & Fischer, M. A meta-analysis of trait differences between invasive and non-invasive plant species. Ecol. Lett. 13, 235–245 (2010).PubMed 
Article 

Google Scholar 
van Kleunen, M., Dawson, W. & Maurel, N. Characteristics of successful alien plants. Mol. Ecol. 24, 1954–1968 (2015).PubMed 
Article 

Google Scholar 
Essl, F. et al. Drivers of the relative richness of naturalized and invasive plant species on Earth. AoB Plants 11, plz051 (2019).PubMed 
PubMed Central 
Article 

Google Scholar 
Winkler, D. E., Gremer, J. R., Chapin, K. J., Kao, M. & Huxman, T. E. Rapid alignment of functional trait variation with locality across the invaded range of Sahara mustard (Brassica tournefortii). Am. J. Bot. 105, 1188–1197 (2018).PubMed 
Article 

Google Scholar 
Divíšek, J. et al. Similarity of introduced plant species to native ones facilitates naturalization, but differences enhance invasion success. Nat. Commun. 9, 4631 (2018).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Banerjee, A. K., Prajapati, J., Bhowmick, A. R., Huang, Y. & Mukherjee, A. Different factors influence naturalization and invasion processes – a case study of Indian alien flora provides management insights. J. Environ. Manag. 294, 113054 (2021).Article 

Google Scholar 
Ni, M. et al. Invasion success and impacts depend on different characteristics in non-native plants. Divers. Distrib. 27, 1194–1207 (2021).Article 

Google Scholar 
Fristoe, T. S. et al. Dimensions of invasiveness: links between local abundance, geographic range size, and habitat breadth in Europe’s alien and native floras. Proc. Natl Acad. Sci. USA 118, e2021173118 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Omer, A. et al. Characteristics of the naturalized flora of Southern Africa largely reflect the non-random introduction of alien species for cultivation. Ecography 44, 1812–1825 (2021).Article 

Google Scholar 
Pyšek, P. et al. Naturalization of central European plants in North America: species traits, habitats, propagule pressure, residence time. Ecology 96, 762–774 (2015).PubMed 
Article 

Google Scholar 
Omer, A., Kordofani, M., Gibreel, H. H., Pyšek, P. & van Kleunen, M. The alien flora of Sudan and South Sudan: taxonomic and biogeographical composition. Biol. Invasions 23, 2033–2045 (2021).Article 

Google Scholar 
Duncan, R. P. & Williams, P. A. Darwin’s naturalization hypothesis challenged. Nature 417, 608–609 (2002).CAS 
PubMed 
Article 

Google Scholar 
Daehler, C. C. Darwin’s naturalization hypothesis revisited. Am. Nat. 158, 324–330 (2001).CAS 
PubMed 
Article 

Google Scholar 
Pyšek, P. Is there a taxonomic pattern to plant invasions? Oikos 82, 282–294 (1998).Article 

Google Scholar 
Tan, J., Pu, Z., Ryberg, W. A. & Jiang, L. Resident–invader phylogenetic relatedness, not resident phylogenetic diversity, controls community invasibility. Am. Nat. 186, 59–71 (2015).PubMed 
Article 

Google Scholar 
Thuiller, W. et al. Resolving Darwin’s naturalization conundrum: a quest for evidence. Divers. Distrib. 16, 461–475 (2010).Article 

Google Scholar 
Loiola, P. P. et al. Invaders among locals: alien species decrease phylogenetic and functional diversity while increasing dissimilarity among native community members. J. Ecol. 106, 2230–2241 (2018).Article 

Google Scholar 
Lososová, Z. et al. Alien plants invade more phylogenetically clustered community types and cause even stronger clustering. Glob. Ecol. Biogeogr. 24, 786–794 (2015).Article 

Google Scholar 
Marx, H. E., Giblin, D. E., Dunwiddie, P. W. & Tank, D. C. Deconstructing Darwin’s naturalization conundrum in the San Juan Islands using community phylogenetics and functional traits. Divers. Distrib. 22, 318–331 (2016).Article 

Google Scholar 
Darwin, C. On the Origin of Species by Means of Natural Selection or the Preservation of Favoured Races in the Struggle for Life (John Murray, 1859).Procheş, Ş., Wilson, J. R. U., Richardson, D. M. & Rejmánek, M. Searching for phylogenetic pattern in biological invasions. Glob. Ecol. Biogeogr. 17, 5–10 (2008).
Google Scholar 
Diez, J. M., Sullivan, J. J., Hulme, P. E., Edwards, G. & Duncan, R. P. Darwin’s naturalization conundrum: dissecting taxonomic patterns of species invasions. Ecol. Lett. 11, 674–681 (2008).PubMed 
Article 

Google Scholar 
Cadotte, M. W., Campbell, S. E., Li, S. P., Sodhi, D. S. & Mandrak, N. E. Preadaptation and naturalization of nonnative species: Darwin’s two fundamental insights into species invasion. Annu Rev. Plant Biol. 69, 661–684 (2018).CAS 
PubMed 
Article 

Google Scholar 
van Kleunen, M., Bossdorf, O. & Dawson, W. The ecology and evolution of alien plants. Annu. Rev. Ecol. Evol. Syst. 49, 25–47 (2018).Article 

Google Scholar 
Park, D. S., Feng, X., Maitner, B. S., Ernst, K. C. & Enquist, B. J. Darwin’s naturalization conundrum can be explained by spatial scale. Proc. Natl Acad. Sci. USA 117, 10904–10910 (2020).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Diez, J. M. et al. Learning from failures: testing broad taxonomic hypotheses about plant naturalization. Ecol. Lett. 12, 1174–1183 (2009).PubMed 
Article 

Google Scholar 
Malecore, E. M., Dawson, W., Kempel, A., Müller, G. & van Kleunen, M. Nonlinear effects of phylogenetic distance on early-stage establishment of experimentally introduced plants in grassland communities. J. Ecol. 107, 781–793 (2019).Article 

Google Scholar 
Schaefer, H., Hardy, O. J., Silva, L., Barraclough, T. G. & Savolainen, V. Testing Darwin’s naturalization hypothesis in the Azores. Ecol. Lett. 14, 389–396 (2011).PubMed 
Article 

Google Scholar 
Strauss, S. Y., Webb, C. O. & Salamin, N. Exotic taxa less related to native species are more invasive. Proc. Natl Acad. Sci. USA 103, 5841–5845 (2006).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Li, S.-p. et al. The effects of phylogenetic relatedness on invasion success and impact: deconstructing Darwin’s naturalisation conundrum. Ecol. Lett. 18, 1285–1292 (2015).PubMed 
Article 

Google Scholar 
Pellock, S., Thompson, A., He, K., Mecklin, C. & Yang, J. Validity of Darwin’s naturalization hypothesis relates to the stages of invasion. Community Ecol. 14, 172–179 (2013).Article 

Google Scholar 
Blackburn, T. M. et al. A proposed unified framework for biological invasions. Trends Ecol. Evol. 26, 333–339 (2011).PubMed 
Article 

Google Scholar 
van Kleunen, M. et al. Economic use of plants is key to their naturalization success. Nat. Commun. 11, 3201 (2020).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Broennimann, O. et al. Distance to native climatic niche margins explains establishment success of alien mammals. Nat. Commun. 12, 2353 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Carboni, M. et al. What it takes to invade grassland ecosystems: traits, introduction history and filtering processes. Ecol. Lett. 19, 219–229 (2016).PubMed 
Article 

Google Scholar 
Milbau, A. & Stout, J. C. Factors associated with alien plants transitioning from casual, to naturalized, to invasive. Conserv. Biol. 22, 308–317 (2008).PubMed 
Article 

Google Scholar 
Dawson, W., Burslem, D. F. R. P. & Hulme, P. E. Factors explaining alien plant invasion success in a tropical ecosystem differ at each stage of invasion. J. Ecol. 97, 657–665 (2009).Article 

Google Scholar 
Rejmánek, M. in Invasive Species and Biodiversity Management (eds Schei, P. J. & Vilken, A.) 79–102 (Kluwer Academic, 1998).Rejmánek, M. A theory of seed plant invasiveness: the first sketch. Biol. Conserv. 78, 171–181 (1996).Article 

Google Scholar 
Maurel, N., Hanspach, J., Kuhn, I., Pysek, P. & van Kleunen, M. Introduction bias affects relationships between the characteristics of ornamental alien plants and their naturalization success. Glob. Ecol. Biogeogr. 25, 1500–1509 (2016).Article 

Google Scholar 
Glen, H. F. Cultivated Plants of Southern Africa: Botanical Names, Common Names, Origins, Literature (National Botanical Institute, 2002).Reichard, S. H. & White, P. Horticulture as a pathway of invasive plant introductions in the United States. Bioscience 51, 103–113 (2001).Article 

Google Scholar 
Faulkner, K. T., Robertson, M. P., Rouget, M. & Wilson, J. R. U. Understanding and managing the introduction pathways of alien taxa: South Africa as a case study. Biol. Invasions 18, 73–87 (2016).Article 

Google Scholar 
Dodd, A. J., Burgman, M. A., McCarthy, M. A. & Ainsworth, N. The changing patterns of plant naturalization in Australia. Divers. Distrib. 21, 1038–1050 (2015).Article 

Google Scholar 
Lambdon, P.-W. et al. Alien flora of Europe: species diversity, temporal trends, geographical patterns and research needs. Preslia 80, 101–149 (2008).
Google Scholar 
Bennett, B. M. Naturalising Australian trees in South Africa: climate, exotics and experimentation. J. South. Afr. Stud. 37, 265–280 (2011).Article 

Google Scholar 
Richardson, D. M. et al. in Biological Invasions in South Africa (eds van Wilgen, B. W. et al.) 67–96 (Springer, 2020).Li, S.-p. et al. Contrasting effects of phylogenetic relatedness on plant invader success in experimental grassland communities. J. Appl. Ecol. 52, 89–99 (2015).CAS 
Article 

Google Scholar 
Duarte, M., Verdú, M., Cavieres, L. A. & Bustamante, R. O. Plant–plant facilitation increases with reduced phylogenetic relatedness along an elevation gradient. Oikos 130, 248–259 (2021).Article 

Google Scholar 
Verdú, M., Rey, P. J., Alcántara, J. M., Siles, G. & Valiente-Banuet, A. Phylogenetic signatures of facilitation and competition in successional communities. J. Ecol. 97, 1171–1180 (2009).Article 

Google Scholar 
Valiente-Banuet, A. & Verdu, M. Plant facilitation and phylogenetics. Annu. Rev. Ecol. Evol. Syst. 44, 347–366 (2013).Article 

Google Scholar 
Anacker, B. L. & Strauss, S. Y. Ecological similarity is related to phylogenetic distance between species in a cross-niche field transplant experiment. Ecology 97, 1807–1818 (2016).PubMed 
Article 

Google Scholar 
Dostál, P. Plant competitive interactions and invasiveness: searching for the effects of phylogenetic relatedness and origin on competition intensity. Am. Nat. 177, 655–667 (2011).PubMed 
Article 

Google Scholar 
Levin, S. C., Crandall, R. M., Pokoski, T., Stein, C. & Knight, T. M. Phylogenetic and functional distinctiveness explain alien plant population responses to competition. Proc. R. Soc. B 287, 20201070 (2020).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Williams, E. W., Zeldin, J., Semski, W. R., Hipp, A. L. & Larkin, D. J. Phylogenetic distance and resource availability mediate direction and strength of plant interactions in a competition experiment. Oecologia 197, 459–469 (2021).PubMed 
Article 

Google Scholar 
Bezeng, S. B., Davies, J. T., Yessoufou, K., Maurin, O. & Van der Bank, M. Revisiting Darwin’s naturalization conundrum: explaining invasion success of non-native trees and shrubs in Southern Africa. J. Ecol. 103, 871–879 (2015).Article 

Google Scholar 
Trotta, L. B., Siders, Z. A., Sessa, E. B. & Baiser, B. The role of phylogenetic scale in Darwin’s naturalization conundrum in the critically imperilled pine rockland ecosystem. Divers. Distrib. 27, 618–631 (2021).Article 

Google Scholar 
Sol, D. et al. A test of Darwin’s naturalization conundrum in birds reveals enhanced invasion success in the presence of close relatives. Ecol. Lett. 25, 661–672 (2022).PubMed 
Article 

Google Scholar 
Smith, S. A. & Brown, J. W. Constructing a broadly inclusive seed plant phylogeny. Am. J. Bot. 105, 302–314 (2018).PubMed 
Article 

Google Scholar 
Henderson, L. Comparisons of invasive plants in Southern Africa originating from southern temperate, northern temperate and tropical regions. Bothalia 36, 201–222 (2006).Article 

Google Scholar 
Cayuela, L., Stein, A. & Oksanen, J. Taxonstand: Taxonomic Standardization of Plant Species Names. R package version 2.2. https://CRAN.R-project.org/package=Taxonstand (R Foundation for Statistical Computing, Vienna, 2019).Weigelt, P., König, C. & Kreft, H. GIFT – A Global Inventory of Floras and Traits for macroecology and biogeography. J. Biogeogr. 47, 16–43 (2020).Article 

Google Scholar 
van Kleunen, M. et al. The Global Naturalized Alien Flora (GloNAF) database. Ecology 100, e02542 (2019).PubMed 
Article 

Google Scholar 
Zengeya, T. A. & Wilson, J. R. (eds) The Status of Biological Invasions and Their Management in South Africa in 2019 (South African National Biodiversity Institute and DSI-NRF Centre of Excellence for Invasion Biology, 2021).Revell, L. J. phytools: an R package for phylogenetic comparative biology (and other things). Methods Ecol. Evol. 3, 217–223 (2012).Article 

Google Scholar 
R: A Language and Environment for Statistical Computing v.3.6.1 (R Foundation for Statistical Computing, 2019).Zuur, A. F., Ieno, E. N., Walker, N. J., Saveliev, A. A. & Smith, G. M. Mixed Effects Models and Extensions in Ecology with R Vol. 574 (Springer, 2009).Schielzeth, H. Simple means to improve the interpretability of regression coefficients. Methods Ecol. Evol. 1, 103–113 (2010).Article 

Google Scholar 
Nagelkerke, N. J. D. A note on a general definition of the coefficient of determination. Biometrika 78, 691–692 (1991).Article 

Google Scholar 
rcompanion: Functions to support extension education program evaluation v. 2.4.1 (R Foundation for Statistical Computing, 2021).Tung Ho, L. S. & Ané, C. A linear-time algorithm for Gaussian and non-Gaussian trait evolution models. Syst. Biol. 63, 397–408 (2014).Article 

Google Scholar  More

Université Paris-Saclay, CNRS, AgroParisTech, Ecologie Systématique Evolution, Orsay, FranceChristophe Diagne, Anne-Charlotte Vaissière & Franck CourchampUnité Biologie des Organismes et Ecosystèmes Aquatiques (BOREA, UMR 7208), Muséum national d’Histoire naturelle, Sorbonne Université, Université de Caen Normandie, CNRS, IRD, Université des Antilles, Paris, FranceBoris LeroyISEM, Univ. Montpellier, CNRS, IRD, Montpellier, FranceRodolphe E. GozlanMIVEGEC, Univ. Montpellier, IRD, CNRS, Montpellier, FranceDavid RoizInstitute of Hydrobiology, Biology Centre of the Czech Academy of Sciences, České Budějovice, Czech RepublicIvan JarićDepartment of Ecosystem Biology, Faculty of Science, University of South Bohemia, České Budějovice, Czech RepublicIvan JarićCEE-M, UMR5211, Univ. Montpellier, CNRS, INRAE, Institut Agro, Montpellier, FranceJean-Michel SallesGlobal Ecology, College of Science and Engineering, Flinders University, Adelaide, South Australia, AustraliaCorey J. A. Bradshaw More

Price, D. T. et al. Anticipating the consequences of climate change for Canada’s boreal forest ecosystems. Environ. Rev. 21, 322–365 (2013).Article 

Google Scholar 
Wang, Y., Hogg, H. E., Price, T. D., Edwards, J. & Williamson, T. Past and projected future changes in moisture conditions in the Canadian boreal forest. Forestry Chron. 90, 678–691 (2014).Article 

Google Scholar 
Piao, S. et al. Plant phenology and global climate change: current progresses and challenges. Glob. Chang. Biol. 25, 1922–1940 (2019).ADS 
MathSciNet 
Article 

Google Scholar 
Lu, P., Parker, W. C., Colombo, S. J. & Skeates, D. A. Temperature-induced growing season drought threatens survival and height growth of white spruce in southern Ontario, Canada. Forest Ecol. Manag. 448, 355–363 (2019).Article 

Google Scholar 
Giorgi, F., Raffaele, F. & Coppola, E. The response of precipitation characteristics to global warming from climate projections. Earth Syst. Dyn. 10, 73–89 (2019).ADS 
Article 

Google Scholar 
Sherwood, S. & Fu, Q. A drier future? Science 343, 737–739 (2014).ADS 
CAS 
Article 

Google Scholar 
Seager, R. et al. Dynamical and thermodynamical causes of large-scale changes in the hydrological cycle over North America in response to global warming. J. Clim. 27, 7921–7948 (2014).ADS 
Article 

Google Scholar 
Tam, B. Y. et al. CMIP5 drought projections in Canada based on the Standardized Precipitation Evapotranspiration Index. Can. Water Resour. J. 44, 90–107 (2019).Article 

Google Scholar 
Wu, Z., Dijkstra, P., Koch, G. W., Peñuelas, J. & Hungate, B. A. Responses of terrestrial ecosystems to temperature and precipitation change: a meta-analysis of experimental manipulation. Glob. Chang. Biol. 17, 927–942 (2011).ADS 
Article 

Google Scholar 
Zhao, J., Hartmann, H., Trumbore, S., Ziegler, W. & Zhang, Y. High temperature causes negative whole-plant carbon balance under mild drought. New Phytol. 200, 330–339 (2013).CAS 
Article 

Google Scholar 
Reich, P. B. et al. Effects of climate warming on photosynthesis in boreal tree species depend on soil moisture. Nature 562, 263–267 (2018).ADS 
CAS 
Article 

Google Scholar 
Hansen, W. D. & Turner, M. G. Origins of abrupt change? Postfire subalpine conifer regeneration declines nonlinearly with warming and drying. Ecol. Monogr. 89, e01340 (2019).Article 

Google Scholar 
Girardin, M. P. et al. No growth stimulation of Canada’s boreal forest under half-century of combined warming and CO2 fertilization. Proc. Natl Acad. Sci. USA 113, E8406–E8414 (2016).CAS 
Article 

Google Scholar 
Sulla-Menashe, D., Woodcock, C. E. & Friedl, M. A. Canadian boreal forest greening and browning trends: an analysis of biogeographic patterns and the relative roles of disturbance versus climate drivers. Environ. Res. Lett. 13, 014007 (2018).ADS 
Article 

Google Scholar 
Peng, C. et al. A drought-induced pervasive increase in tree mortality across Canada’s boreal forests. Nat. Clim. Chang. 1, 467–471 (2011).ADS 
Article 

Google Scholar 
Ma, Z. et al. Regional drought-induced reduction in the biomass carbon sink of Canada’s boreal forests. Proc. Natl Acad. Sci. USA 109, 2423–2427 (2012).ADS 
CAS 
Article 

Google Scholar 
Ju, J. & Masek, J. G. The vegetation greenness trend in Canada and US Alaska from 1984–2012 Landsat data. Remote Sens. Environ. 176, 1–16 (2016).ADS 
Article 

Google Scholar 
D’Orangeville, L. et al. Beneficial effects of climate warming on boreal tree growth may be transitory. Nat. Commun. 9, 3213 (2018).ADS 
Article 

Google Scholar 
Johnstone, J. F. et al. Changing disturbance regimes, ecological memory and forest resilience. Front. Ecol. Environ. 14, 369–378 (2016).Article 

Google Scholar 
Rodgers, V. L., Smith, N. G., Hoeppner, S. S. & Dukes, J. S. Warming increases the sensitivity of seedling growth capacity to rainfall in six temperate deciduous tree species. AoB Plants 10, ply003 (2018).Article 

Google Scholar 
Moyes, A. B., Castanha, C., Germino, M. J. & Kueppers, L. M. Warming and the dependence of limber pine (Pinus flexilis) establishment on summer soil moisture within and above its current elevation range. Oecologia 171, 271–282 (2013).ADS 
Article 

Google Scholar 
Balducci, L. et al. How do drought and warming influence survival and wood traits of Picea mariana saplings? J. Exp. Bot. 66, 377–389 (2015).CAS 
Article 

Google Scholar 
Reich, P. B. et al. Geographic range predicts photosynthetic and growth response to warming in co-occurring tree species. Nat. Clim. Chang. 5, 148–152 (2015).ADS 
CAS 
Article 

Google Scholar 
Coursolle, C. et al. Moving towards carbon neutrality: CO2 exchange of a black spruce forest ecosystem during the first 10 years of recovery after harvest. Can. J. Forest Res. 42, 1908–1918 (2012).CAS 
Article 

Google Scholar 
Khomik, M., Williams, C. A., Vanderhoof, M. K., MacLean, R. G. & Dillen, S. Y. On the causes of rising gross ecosystem productivity in a regenerating clearcut environment: leaf area vs. species composition. Tree Physiol. 34, 686–700 (2014).Article 

Google Scholar 
Engelbrecht, B. et al. Drought sensitivity shapes species distribution patterns in tropical forests. Nature 447, 80–82 (2007).ADS 
CAS 
Article 

Google Scholar 
Friedman, S. K. & Reich, P. B. Regional legacies of logging: departure from presettlement forest conditions in northern Minnesota. Ecol. Appl. 15, 726–744 (2005).Article 

Google Scholar 
Burrill, E. A. et al. The Forest Inventory and Analysis Database: Database Description and User Guide Version 9.0.1 for Phase 2 https://www.fia.fs.fed.us/library/database-documentation/ (Forest Service, US Department of Agriculture, 2022).Cumming, S. G. et al. A gap analysis of tree species representation in the protected areas of the Canadian boreal forest: applying a new assemblage of digital Forest Resource Inventory data. Can. J. Forest Res. 45, 163–173 (2015).Article 

Google Scholar 
Brook, B. W., Ellis, E. C., Perring, M. P., Mackay, A. W. & Blomqvist, L. Does the terrestrial biosphere have planetary tipping points? Trends Ecol. Evol. 28, 396–401 (2013).Article 

Google Scholar 
Reyer, C. P. O. et al. Forest resilience and tipping points at different spatio-temporal scales: approaches and challenges. J. Ecol. 103, 5–15 (2015).ADS 
Article 

Google Scholar 
Stralberg, D. et al. Climate‐change refugia in boreal North America: what, where, and for how long? Front. Ecol. Environ. 18, 261–270 (2020).Article 

Google Scholar 
Etterson, J. R., Cornett, M. W., White, M. A. & Kavajecz, L. C. Assisted migration across fixed seed zones detects adaptation lags in two major North American tree species. Ecol. Appl. 30, e02092 (2020).Article 

Google Scholar 
Solarik, K. A., Cazelles, K., Messier, C., Bergeron, Y. & Gravel, D. Priority effects will impede range shifts of temperate tree species into the boreal forest. J. Ecol. 108, 1155–1173 (2020).Article 

Google Scholar 
Stefanski, A., Bermudez, R., Sendall, K. M., Montgomery, R. A. & Reich, P. B. Surprising lack of sensitivity of biochemical limitation of photosynthesis of nine tree species to open‐air experimental warming and reduced rainfall in a southern boreal forest. Glob. Chang. Biol. 26, 746–759 (2020).ADS 
Article 

Google Scholar 
Perala, D. A. How endemic injuries affect early growth of aspen suckers. Can. J. Forest Res. 14, 755–762 (1984).Article 

Google Scholar 
Buckman, R. E. Effects of prescribed burning on hazel in Minnesota. Ecology 45, 626–629 (1964).Article 

Google Scholar 
Harvey, B. D. & Bergeron, Y. Site patterns of natural regeneration following clear-cutting in northwestern Quebec. Can. J. Forest Res. 19, 1458–1469 (1989).Article 

Google Scholar 
Harris, I. et al. Version 4 of the CRU TS monthly high-resolution gridded multivariate climate dataset. Sci. Data 7, 109 (2020).Article 

Google Scholar 
Peters, M. P., Prasad, A. M., Matthews, S. N. & Iverson, L. R. Climate Change Tree Atlas, Version 4 https://www.nrs.fs.fed.us/atlas (Northern Research Station and Northern Institute of Applied Climate Science, US Forest Service, 2020)Niinemets, Ü. & Valladares, F. Tolerance to shade, drought, and waterlogging of temperate Northern Hemisphere trees and shrubs. Ecol. Monogr. 76, 521–547 (2006).Article 

Google Scholar  More

Short-term experimentsPotential toxic and cryoprotection effects of different CPA combinationsFocusing on toxicity bioassays (Figs. 1A, 2A), although there were certain CPA combinations that yielded significant abnormality percentages compared to controls, in general the CPA combinations did not yield any significant toxic effect. The use of Milli-Q Water instead of FSW did not enhance normal larval development after CPA exposure, neither did the addition of PVP at the concentrations tested, even in combination with trehalose (TRE) (p  > 0.05). In fact, the highest concentrations of PVP used in this experiment (9 and 12%) yielded significant abnormal development on exposed trochophores (Fig. 1A) (p  More

Field CB, Behrenfeld MJ, Randerson JT, Falkowski P. Primary production of the biosphere: integrating terrestrial and oceanic components. Science. 1998;281:237–40. https://doi.org/10.1126/science.281.5374.237CAS 
Article 
PubMed 

Google Scholar 
Falkowski PG, Fenchel T, Delong EF. The microbial engines that drive Earth’s biogeochemical cycles. Science. 2008;320:1034–9. https://doi.org/10.1126/science.1153213CAS 
Article 
PubMed 

Google Scholar 
Gattuso J-P, Magnan A, Billé R, Cheung WWL, Howes EL, Joos F, et al. Contrasting futures for ocean and society from different anthropogenic CO2 emissions scenarios. Science. 2015;349:aac4722. https://doi.org/10.1126/science.aac4722Steinacher M, Joos F, Frölicher TL, Bopp L, Cadule P, Cocco V, et al. Projected 21st century decrease in marine productivity: a multi-model analysis. Biogeosciences. 2010;7:979–1005. https://doi.org/10.5194/bg-7-979-2010CAS 
Article 

Google Scholar 
Henson SA, Cael BB, Allen SR, Dutkiewicz S. Future phytoplankton diversity in a changing climate. Nat Commun. 2021;12:5372. https://doi.org/10.1038/s41467-021-25699-wCAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Thomas MK, Kremer CT, Klausmeier CA, Litchman E. A global pattern of thermal adaptation in marine phytoplankton. Science. 2012;338:1085–8. https://doi.org/10.1126/science.1224836CAS 
Article 
PubMed 

Google Scholar 
Collins S, Boyd PW, Doblin MA. Evolution, microbes, and changing ocean conditions. Annu Rev Mar Sci. 2020;12:181–208. https://doi.org/10.1146/annurev-marine-010318-095311Article 

Google Scholar 
Schaum CE, Buckling A, Smirnoff N, Studholme DJ, Yvon-Durocher G. Environmental fluctuations accelerate molecular evolution of thermal tolerance in a marine diatom. Nat Commun. 2018;9:1719. https://doi.org/10.1038/s41467-018-03906-5CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Lohbeck KT, Riebesell U, Reusch TBH. Adaptive evolution of a key phytoplankton species to ocean acidification. Nat Geosci. 2012;5:346–51. https://doi.org/10.1038/ngeo1441CAS 
Article 

Google Scholar 
Jin P, Gao K, Beardall J. Evolutionary responses of a coccolithophorid Gephyrocapsa oceanica to ocean acidification. Evolution. 2013;67:1869–78. https://doi.org/10.1111/evo.12112CAS 
Article 
PubMed 

Google Scholar 
Schlüter L, Lohbeck KT, Gutowska MA, Gröger JP, Riebesell U, Reusch TBH. Adaptation of a globally important coccolithophore to ocean warming and acidification. Nat Clim Change. 2014;4:1024–30. https://doi.org/10.1038/nclimate2379CAS 
Article 

Google Scholar 
Listmann L, LeRoch M, Schlüter L, Thomas MK, Reusch TBH. Swift thermal reaction norm evolution in a key marine phytoplankton species. Evol Appl. 2016;9:1156–64. https://doi.org/10.1111/eva.12362Article 
PubMed 
PubMed Central 

Google Scholar 
Zhong J, Guo Y, Liang Z, Huang Q, Lu H, Pan J, et al. Adaptation of a marine diatom to ocean acidification and warming reveals constraints and trade-offs. Sci Total Environ. 2021;771:145167. https://doi.org/10.1016/j.scitotenv.2021.145167CAS 
Article 
PubMed 

Google Scholar 
Brennan GL, Colegrave N, Collins S. Evolutionary consequences of multidriver environmental change in an aquatic primary producer. Proc Natl Acad Sci USA. 2017;114:9930–5. https://doi.org/10.1073/pnas.1703375114CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Zhang S, Wu Y, Lin L, Wang D. Molecular insights into the circadian clock in marine diatoms. Acta Oceano Sin. 2022;41:1–12. https://doi.org/10.1007/s13131-021-1962-4Article 

Google Scholar 
Nagelkerken I, Connell SD. Global alteration of ocean ecosystem functioning due to increasing human CO2 emissions. Proc Natl Acad Sci USA. 2015;112:13272–7. https://doi.org/10.1073/pnas.1510856112CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Boyd PW, Collins S, Dupont S, Fabricius K, Gattuso JP, Havenhand J, et al. Experimental strategies to assess the biological ramifications of multiple drivers of global ocean change-a review. Glob Change Biol. 2018;24:2239–61. https://doi.org/10.1111/gcb.14102Article 

Google Scholar 
Matsuda Y, Nakajima K, Tachibana M. Recent progresses on the genetic basis of the regulation of CO2 acquisition systems in response to CO2 concentration. Photosynth Res. 2011;109:191–203. https://doi.org/10.1007/s11120-011-9623-7CAS 
Article 
PubMed 

Google Scholar 
Ohno N, Inoue T, Yamashiki R, Nakajima K, Kitahara Y, Ishibashi M, et al. CO2-cAMP-responsive cis-elements targeted by a transcription factor with CREB/ATF-like basic zipper domain in the marine diatom Phaeodactylum tricornutum. Plant Physiol. 2012;158:499–513. https://doi.org/10.1104/pp.111.190249CAS 
Article 
PubMed 

Google Scholar 
Hennon GMM, Ashworth J, Groussman RD, Berthiaume C, Morales RL, Baliga NS, et al. Diatom acclimation to elevated CO2 via cAMP signalling and coordinated gene expression. Nat Clim Change. 2015;5:761–5. https://doi.org/10.1038/nclimate2683CAS 
Article 

Google Scholar 
Toseland A, Daines SJ, Clark JR, Kirkham A, Strauss J, Uhlig C, et al. The impact of temperature on marine phytoplankton resource allocation and metabolism. Nat Clim Change. 2013;3:979–84. https://doi.org/10.1038/nclimate1989CAS 
Article 

Google Scholar 
Gao K, Beardall J, Häder DP, Hall-Spencer JM, Gao G, Hutchins DA. Effects of ocean acidification on marine photosynthetic organisms under the concurrent influences of warming, UV radiation, and deoxygenation. Front Mar Sci. 2019;6:322. https://doi.org/10.3389/fmars.2019.00322Article 

Google Scholar 
Tu L, Su P, Zhang Z, Gao L, Wang J, Hu T, et al. Genome of Tripterygium wilfordii and identification of cytochrome P450 involved in triptolide biosynthesis. Nat Commun. 2020;11:971. https://doi.org/10.1038/s41467-020-14776-1CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Treves H, Siemiatkowska B, Luzarowska U, Murik O, Fernandez-Pozo N, Moraes TA, et al. Multi-omics reveals mechanisms of total resistance to extreme illumination of a desert alga. Nat Plants. 2020;6:1031–43. https://doi.org/10.1038/s41477-020-0729-9CAS 
Article 
PubMed 

Google Scholar 
Van den Bergh B, Swings T, Fauvart M, Michels J. Experimental design, population dynamics, and diversity in microbial experimental evolution. Microbiol Mol Biol Rev. 2018;82:e00008–18.PubMed 
PubMed Central 

Google Scholar 
Elena SF, Lenski RE. Evolution experiments with microorganisms: the dynamics and genetic bases of adaptation. Nat Rev Genet. 2003;4:457–69. https://doi.org/10.1038/nrg1088CAS 
Article 
PubMed 

Google Scholar 
Colegrave N, Collins S. Experimental evolution: experimental evolution and evolvability. Heredity. 2008;100:464–70. https://doi.org/10.1038/sj.hdy.6801095CAS 
Article 
PubMed 

Google Scholar 
Jin P, Ji Y, Huang Q, Li P, Pan J, Lu H, et al. A reduction in metabolism explains the trade‐offs associated with the long‐term adaptation of phytoplankton to high CO2 concentrations. N Phytol. 2022;233:2155–67. https://doi.org/10.1111/nph.17917CAS 
Article 

Google Scholar 
Flombaum P, Gallegos JL, Gordillo RA, Rincón J, Zabala LL, Jiao N, et al. Present and future global distributions of the marine Cyanobacteria Prochlorococcus and Synechococcus. Proc Natl Acad Sci USA. 2013;110:9824–9. https://doi.org/10.1073/pnas.1307701110CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Hutchins DA, Walworth NG, Webb EA, Saito MA, Moran D, Mcllvin MR, et al. Irreversibly increased nitrogen fixation in Trichodesmium experimentally adapted to elevated carbon dioxide. Nat Commun. 2015;6:8155. https://doi.org/10.1038/ncomms9155Article 
PubMed 

Google Scholar 
Padfield D, Yvon-Durocher G, Buckling A, Jennings S, Yvon-Durocher G. Rapid evolution of metabolic traits explains thermal adaptation in phytoplankton. Ecol Lett. 2016;19:133–42.Article 

Google Scholar 
Coles VJ, Stukel MR, Brooks MT, Burd A, Crump BC, Moran MA, et al. Ocean biogeochemistry modeled with emergent trait-based genomics. Science. 2017;358:1149–54. https://doi.org/10.1126/science.aan5712CAS 
Article 
PubMed 

Google Scholar 
Linnen CR, Kingsley EP, Jensen JD, Hoekstra HE. On the origin and spread of an adaptive allele in deer mice. Science. 2009;325:1095–8. https://doi.org/10.1126/science.1175826CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Van’t Hof AE, Campagne P, Rigden DJ, Yung CJ, Lingley J, Quail MA, et al. The industrial melanism mutation in British peppered moths is a transposable element. Nature. 2016;534:102–5. https://doi.org/10.1038/nature17951CAS 
Article 
PubMed 

Google Scholar 
Bitter MC, Kapsenberg L, Gattuso JP, Pfister CA. Standing genetic variation fuels rapid adaptation to ocean acidification. Nat Commun. 2019;10:5821. https://doi.org/10.1038/s41467-019-13767-1CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Lai YT, Yeung CK, Omland KE, Pang EL, Hao Y, Liao BY, et al. Standing genetic variation as the predominant source for adaptation of a songbird. Proc Natl Acad Sci USA. 2019;116:2152–7. https://doi.org/10.1073/pnas.1813597116Armbrust EV. The life of diatoms in the world’s oceans. Nature. 2009;459:185–92. https://doi.org/10.1038/nature08057CAS 
Article 
PubMed 

Google Scholar 
Rastogi A, Vieira FRJ, Deton-Cabanillas AF, Veluchamy A, Cantrel C, Wang G, et al. A genomics approach reveals the global genetic polymorphism, structure, and functional diversity of ten accessions of the marine model diatom Phaeodactylum tricornutum. ISME J. 2020;14:347–63. https://doi.org/10.1038/s41396-019-0528-3Article 
PubMed 

Google Scholar 
Jin P, Agustí S. Fast adaptation of tropical diatoms to increased warming with trade-offs. Sci Rep. 2018;8:17771. https://doi.org/10.1038/s41598-018-36091-yCAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Barton S, Jenkins J, Buckling A, Schaum CE, Smirnoff N, Raven JA, et al. Evolutionary temperature compensation of carbon fixation in marine phytoplankton. Ecol Lett. 2020;23:722–33.Article 

Google Scholar 
Guillard RR, Ryther JH. Studies of marine planktonic diatoms: I. Cyclotella nana Hustedt, and Detonula confervacea (Cleve) Gran. Can J Microbiol. 1962;8:229–39. https://doi.org/10.1139/m62-029CAS 
Article 
PubMed 

Google Scholar 
Huysman MJ, Martens C, Vandepoele K, Gillard J, Rayko E, Heijde M, et al. Genome-wide analysis of the diatom cell cycle unveils a novel type of cyclins involved in environmental signaling. Genome Biol. 2010;11:R17. https://doi.org/10.1186/gb-2010-11-2-r17CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
IPCC. Summary for policymakers. In: Masson-Delmotte V, Zhai P, Pirani A, Connors SL, Péan C, Berger S, et al. editors. Climate change 2021: the physical science basis. Contribution of Working Group I to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Switzerland: IPCC; 2021.Jiang H, Gao K. Effects of lowering temperature during culture on the production of polyunsaturated fatty acids in the marine diatom Phaeodactylum tricornutum (Bacillariophyceae). J Phycol. 2004;40:651–4. https://doi.org/10.1111/j.1529-8817.2004.03112.xCAS 
Article 

Google Scholar 
Pérez EB, Pina IC, Rodríguez LP. Kinetic model for growth of Phaeodactylum tricornutum in intensive culture photobioreactor. Biochem Eng J. 2008;40:520–5. https://doi.org/10.1016/j.bej.2008.02.007CAS 
Article 

Google Scholar 
Boyd PW, Rynearson TA, Armstrong EA, Fu F, Hayashi K, Hu Z, et al. Marine phytoplankton temperature versus growth responses from polar to tropical waters-outcome of a scientific community-wide study. PLoS One. 2013;8:e63091 https://doi.org/10.1371/journal.pone.0063091CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Zeng X, Jin P, Jiang Y, Yang H, Zhong J, Liang Z, et al. Light alters the responses of two marine diatoms to increased warming. Mar Environ Res. 2020;154:104871. https://doi.org/10.1016/j.marenvres.2019.104871CAS 
Article 
PubMed 

Google Scholar 
Chen S, Zhou Y, Chen Y, Gu J. fastp: an ultra-fast all-in-one FASTQ preprocessor. Bioinformatics. 2018;34:i884–i890. https://doi.org/10.1093/bioinformatics/bty560CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Bowler C, Allen AE, Badger JH, Grimwood J, Jabbari K, Kuo A, et al. The Phaeodactylum genome reveals the evolutionary history of diatom genomes. Nature. 2008;456:239–44.CAS 
Article 

Google Scholar 
Li H, Durbin R. Fast and accurate short read alignment with Burrows-Wheeler transform. Bioinformatics. 2009;25:1754–60. https://doi.org/10.1093/bioinformatics/btp324CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Wang K, Li M, Hakonarson H. ANNOVAR: functional annotation of genetic variants from high-throughput sequencing data. Nucleic Acids Res. 2010;38:e164. https://doi.org/10.1093/nar/gkq603CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Langmead B, Salzberg SL. Fast gapped-read alignment with Bowtie 2. Nat Methods. 2012;9:357–9. https://doi.org/10.1038/nmeth.1923CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Kim D, Langmead B, Salzberg SL. HISAT: a fast spliced aligner with low memory requirements. Nat Methods. 2015;12:357–60. https://doi.org/10.1038/nmeth.3317CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Pertea M, Pertea GM, Antonescu CM, Chang TC, Mendell JT, Salzberg SL. StringTie enables improved reconstruction of a transcriptome from RNA-seq reads. Nat Biotechnol. 2015;33:290–5. https://doi.org/10.1038/nbt.3122CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Pertea M, Kim D, Pertea GM, Leek JT, Salzberg SL. Transcript-level expression analysis of RNA-seq experiments with HISAT, StringTie and Ballgown. Nat Protoc. 2016;11:1650–67. https://doi.org/10.1038/nprot.2016.095CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Love MI, Huber W, Anders S. Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2. Genome Biol. 2014;15:550. https://doi.org/10.1186/s13059-014-0550-8CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Gifford RM. Plant respiration in productivity models: conceptualisation, representation and issues for global terrestrial carbon-cycle research. Funct Plant Biol. 2003;30:171–86. https://doi.org/10.1071/FP02083Article 
PubMed 

Google Scholar 
Jassby AD, Platt T. Mathematical formulation of the relationship between photosynthesis and light for phytoplankton. Limnol Oceanogr. 1976;21:540–7. https://doi.org/10.4319/lo.1976.21.4.0540CAS 
Article 

Google Scholar  More

Dow, C. et al. Nature 608, 552–557 (2022).Article 

Google Scholar 
Friedlingstein, P. et al. Earth Syst. Sci. Data 12, 3269–3340 (2020).Article 

Google Scholar 
Menzel, A. & Fabian, P. Nature 397, 659 (1999).Article 

Google Scholar 
Piao, S. et al. Nature Rev. Earth Environ. 1, 14–27 (2020).Article 

Google Scholar 
Cuny, H. E. et al. Nature Plants 1, 15160 (2015).PubMed 
Article 

Google Scholar 
Körner, C. Curr. Opin. Plant Biol. 25, 107–114 (2015).PubMed 
Article 

Google Scholar 
Gessler, A. & Treydte, K. New Phytol. 209, 1338–1340 (2016).PubMed 
Article 

Google Scholar 
Hilty, J., Muller, B., Pantin, F. & Leuzinger, S. New Phytol. 232, 25–41 (2021).PubMed 
Article 

Google Scholar 
Jiang, M. et al. Nature 580, 227–231 (2020).PubMed 
Article 

Google Scholar 
Guillemot, J. et al. New Phytol. 214, 180–193 (2017).PubMed 
Article 

Google Scholar 
Fatichi, S., Pappas, C., Zscheischler, J. & Leuzinger, S. New Phytol. 221, 652–668 (2019).PubMed 
Article 

Google Scholar 
Friend, A. D. et al. Annu. For. Sci. 76, 49 (2019).Article 

Google Scholar 
Zuidema, P. A., Poulter, B. & Frank, D. C. Trends Plant Sci. 23, 1006–1015 (2018).PubMed 
Article 

Google Scholar 
Martínez-Sancho, E., Treydte, K., Lehmann, M. M., Rigling, A. & Fonti, P. New Phytol. https://doi.org/10.1111/nph.18224 (2022).Article 

Google Scholar  More