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3/26/17 Biogeography expands: • Phylogeography (done) PhyloEcoBiogeography: Biome Relationships Jeannine Cavender-Bares et al. 2009 • PhyloEcoBiogeography PhyloEcoBiogeography: Biome Relationships Phylogenetics can inform ecological processes at many scales PhyloEcoBiogeography: Biome Relationships Examined speciation events within Southern Hemisphere continental biome types Examined speciation events within Southern Hemisphere continental biome types 1. Most speciation events of trees (and herbs) occur within same biome type OR between similar biome types 2. Most transoceanic colonizations occur within same biome type 3. Niche conservatism NOT adaptive radiation is seen in S. Hemisphere diversification Michael Crisp et al. (2009) Nature Michael Crisp et al. (2009) Nature Only 356 shifts occurred in 10,800 speciation events within same biome between two biomes 1 3/26/17 PhyloEcoBiogeography: Biome Relationships • Most species are of recent origin • Speciation events involve daughter species in similar communities in similar geographical area — niche conservatism Examined phylogenetic and biogeographical relationships within Seasonally Dry Tropical Forests Toby Pennington et al. (2009) Ann Rev Ecol Syst Toby Pennington et al. (2009) Ann Rev Ecol Syst Chronogram of legume tribe PhyloEcoBiogeography: Biome Relationships PhyloEcoBiogeography: Evolution of Niche • Climatic niche evolution in California Clarkia • Strong correlation of geographic distance and phylogenetic distance! • Do related species share similar climatic/elevation niches? (niche conservatism) • Do related species show significant disparity in climatic/elevation niches? (adaptive radiation) Toby Pennington et al. (2009) Ann Rev Ecol Syst Mantel test 2 3/26/17 PhyloEcoBiogeography: Evolution of Niche Chronogram of 35 diploid Clarkia species C. breweri C. concinna 10mya PhyloEcoBiogeography: Evolution of Niche Mean annual temp Mean temp coldest quarter Mean diurnal temp range Annual precipitation • Temperature and elevation variables show more disparity among close relatives in speciation - adaptive radiation Isothermality Precipitation wettest month Temperature seasonality Precipitation driest month Max temp warmest month Precipitation seasonality Min temp coldest month Precipitation wettest quarter Temp annual range Precipitation driest quarter Mean temp wettest quarter Precipitation warmest quarter Mean temp driest quarter Precipitation coldest quarter Mean temp warmest quarter Elevation • Precipitation shows phylogenetic conservatism – close species are more similar in precipitation niche Downloaded from from rstb.royalsocietypublishing.org rstb.royalsocietypublishing.org on on October October 22, 22, 2010 2010 Downloaded (a) (a) pre-warming pre-warming • Temperature shows significant disparity through time < 5mya - adaptive radiation others, which which has has been been shown shown to to result result in in ecological ecological others, mismatches that that can can have have negative negative fitness fitness conseconsemismatches quences (Stenseth (Stenseth & & Mysterud Mysterud 2002; 2002; Stenseth Stenseth et et al. al. quences 2002; Visser Visser & & Both Both 2005; 2005; Both Both et et al. al. 2006). 2006). Under Under 2002; these circumstances, circumstances, not not only only is is the the inclusion inclusion of of phylophylothese geny an an important important analytical analytical tool tool for for more more properly properly geny assessing statistical statistical trait trait correlations, correlations, but but itit is is vital vital for for assessing understanding the the pattern pattern of of differential differential species’ species’ understanding response to to climate climate change. change. response Willis et et al. al. (2008, (2008, 2009) 2009) recently recently demonstrated demonstrated the the Willis value of of applying applying aa phylogenetic phylogenetic approach approach to to examine examine value the impacts impacts of of climate climate change change using using aa long-term long-term the phenological dataset dataset from from Concord, Concord, MA MA (USA) (USA) that that phenological was initiated initiated by by the the naturalist naturalist and and conservationist conservationist was Henry David David Thoreau Thoreau (Miller-Rushing (Miller-Rushing & & Primack Primack Henry 2008; Primack et al. 2009). They analysed these floris2008; Primack et al. 2009). They analysed these floristic data data using using statistical statistical methods methods that that incorporate incorporate tic phylogenetic history history and and discovered discovered that that clades clades that that phylogenetic have been been less less able able to to respond respond to to climate climate change change by by have adjusting their flowering time have significantly adjusting their flowering time have significantly declined in in abundance. abundance. These These results results can can help help predict predict declined which species species face face aa greater greater risk risk of of regional regional extinction extinction which as climate climate change change is is exacerbated. exacerbated. For For example, example, they they as indicate that that we we should should be be particularly particularly concerned concerned indicate about the the continued continued regional regional loss loss of of species species in in the the about Liliaceae and and Orchidaceae Orchidaceae clades, clades, but but perhaps perhaps less less so so Liliaceae of species species in in the the Brassicaceae Brassicaceae and and Fabaceae. Fabaceae. The The of latter two clades contain species that have been far latter two clades contain species that have been far better able able to to adjust adjust their their phenology phenology to to climate climate better change, and and contain contain fewer fewer species species that that have have declined declined change, in abundance. abundance. However, However, the the extent extent to to which which these these in regional results results are are more more broadly broadly applicable applicable to to other other regional geographically disjunct disjunct plant plant communities, communities, or or to to other other geographically groups of of species species in in the the Tree Tree of of Life, Life, remains remains unexaunexagroups mined. In In particular, particular, do do closely closely related related species species in in mined. different geographical geographical regions, regions, and and in in different different parts parts different of the the Tree Tree of of Life, Life, share share similar similar phenological phenological responses responses of to climate climate change? change? And And is is phenological phenological response response simisimito larly associated associated with with changes changes in in species’ species’ abundance? abundance? larly To address address these these questions, questions, we we extend extend the the approach approach To of Willis Willis et et al. al. (2008) (2008) to to test test for for similar similar trends trends in in plant plant of and bird bird communities communities in in the the United United States States and and the the and UK. First, First, we we compare compare the the phylogenetic phylogenetic distribution distribution UK. of phenological phenological response response traits traits between between the the geographicgeographicof ally disjunct disjunct temperate temperate plant plant communities communities of of Concord, Concord, ally USA and and Chinnor, Chinnor, UK. UK. Second, Second, we we examine examine how how USA phenological response response traits traits contribute contribute to to the the phylogephylogephenological netic pattern pattern of of non-native non-native plant plant species’ species’ success success netic within each each of of these these communities. communities. And And third, third, we we test test within for the the influence influence of of climate climate change change on on the the pattern pattern of of for Consequences of global warming? do all species have ability to track climate change to their species niche? • if different lineages of plants and animals have different adaptations to temperature . . . (b) (b) post-warming post-warming mean mean temperature temperature • Precipitation shows no significant disparity through time - phylogenetic conservatism PhyloEcoBiogeography: is especially especially relevant relevant because because climate climate change change has has is Climate Change greatly altered altered the phenology phenology of of some some species species but but not not greatly the mean mean temperature temperature PhyloEcoBiogeography: Evolution of Niche 3202 C. C. C. C. Davis Davis et et al. al. Phylogeny Phylogeny and and phenology phenology 3202 extinct extinct • then there may be clade specific extinction with global warming Davis et al. 2010 – Importance of phylogeny to the study of phenological response to global climate change Figure 1. 1. Phylogenetic Phylogenetic bias bias in in the the pattern pattern of of species species decline decline Figure owing to to climate climate change. change. A A hypothetical hypothetical phylogeny phylogeny of of organorganowing isms is is depicted depicted during during prepre- and and post-warming post-warming intervals. intervals. The The isms red vertical vertical bars bars over over each each species species represent represent aa climatically climatically relrelred evant trait trait that that is is linked linked to to species species success success (e.g. (e.g. ‘species ‘species evant thermal tolerance’). tolerance’). This This trait trait exhibits exhibits phylogenetic phylogenetic thermal signal—i.e. closely closely related related species species share share similar similar thermal thermal toltolsignal—i.e. erances. An An environmental environmental temperature temperature regime, regime, mean mean erances. annual temperature, temperature, is is depicted depicted by by the the yellow yellow shading shading in in annual the background. background. Following Following warming, warming, some some clades clades of of species species the remain within within their their thermal thermal tolerance tolerance range, range, whereas whereas other other remain clades are are now now well well outside outside their their range range and and become become locally locally clades extirpated. This This results results in in aa phylogenetically phylogenetically biased biased pattern pattern extirpated. of species species loss loss (i.e. (i.e. phylogenetic phylogenetic selectivity). selectivity). of 3 3/26/17 PhyloEcoBiogeography: Climate Change PhyloEcoBiogeography: Climate Change wild geranium Consequences of global warming? do all species have ability to track climate change to their species niche? Downloaded from rstb.royalsocietypublishing.org on October 22, 2010 • used flowering phenology (& migratory bird arrival) data from Thoreau’s Woods in Concord, MA from 1851-2010 3206 C. C. Davis et al. Phylogeny and phenology (a) Chinnor, UK 2 Orchidaceae 1 Araili-/Apiaceae e.g. Daucus 22 Dipsacales p.p. e.g. Ranunculus e.g. Campanulaa e.g. Acer, Aseculus 3 2 e.g. Arabidopsis, Brassica 22 Asterales 18 4 19 e.g. Campanula, Circisum Thoreau Woods 1851 - 2010 4 Brassicaceae 20 Asteralaceae p.p. 19 e.g. Solidago red-eyed vireo 3 Sapindaceae 1 e.g. Scabiosa, Vallerianella 21 Camanulaceae e.g. Ophrys Ranunculaceae p.p. 5 Onagraceae 5 e.g. Oenothera, Epilobium 6 Rosaceae p.p. e.g. Prunus, Pyrus 6 18 7 Rosaceae p.p. e.g. Geum, Rosa 7 8 8 Betulaceae e.g. Betula, Corylus 17 species from the Concord flora depicting changes in abundance Fig. 1. Composite phylogeny of 429 flowering plant from 1900 to 2007. Change in abundance 9 ranged on an integer scale from Boraginaceae !5 to "4, and 17 was calculated as the difference in abundance for each taxon in 1900 and 2007 based on 7 abundance categories Myosotis 9 Fabaceae (0 to 6; see Materials and Methods).e.g.Branch color indicates parsimony character state reconstruction of change in abundance. For simplicity, we have indicated 16 e.g. Lotus, Trifolium this reconstruction by using 4 colors: Rubiaceae red (major 16 decline, !5 to !3), 14 pink (moderate decline, !2), gray (little to no change, !1 to "1), and blue (increase, "2 to 10 e.g. Galium Chinnor versus Concord "4). For the complete character reconstruction and taxon for all internal nodes as the mean 15 labels see Fig. S1. Average decline in abundance was calculated (c) 13 ascertained from 10 change in abundance of descendant nodes weighted with branch length information divergence time estimates. An average decline of 2.5 Time or Tracking Euphorbiaceae Rel. Flowering e.g.using Euphorbia Plantaginaceace p.p.50% 15 or greater, based on our most conservative scoring greater corresponds to a decline in abundance of 6 abundance categories (0 to 5; see0Materials 0.5 1.0 1.5 11 e.g. Veronica in Methods). Clades exhibiting these major declines are indicated with black dots. Each of the most inclusive clades exhibiting these declines are indicated in pink Chinnor/ Lamiales 14 12 Concord and referenced numerically to their clade name.Verbascum Subclades in major decline that are nested within more widely recognized clades are labeled with the more e.g. Mentha, 11 Saxifragales p.p. Rubiaceae/ familiar name followed by pro parte (p.p.). These clades include some of the most charismatic wildflower as anemones and e.g. Ribes, Saxifraga species in New England, such Rubiaceae Caryophyllaceae p.p. bedstraws and bluets (Rubiaceae p.p.), bladderworts buttercups (Ranunculaceae p.p.), asters, campanulas, goldenrods, pussytoes, and12thistles (Asterales), Rosaceae p.p. (Prunus, Malus/Sorbus)/ e.g. Stellaria (Lentibulariaceae), dogwoods (Cornaceae), lilies (Liliales), louseworts and Indian paintbrushes (Orobanchaceae), mints (Lamiaceae p.p.), orchids (Orchidaceae), Rosaceae p.p. (Prunus, Amelanchier) 13 Caryophyllales Rosaceae p.p. (Geum, Rosa)/ primroses (Onograceae p.p.), roses (Rosaceae p.p.), saxifrages (Saxifragales), Indian pipes (Ericales violets (Malpighiales). e.g. Silene, Rumexp.p.), and St. John’s worts and *** PhyloEcoBiogeography: Climate Change Rosaceae p.p. (Rosa, Potentilla) Ranunculaceae p.p. (Ranuculus)/ Ranunculaceae p.p. (Ranunculus) Ranunculaceae (Clematis, Ranuculus)/ Ranunculaceae e.g. Ranunculus 3 Ranuculaceae (Clematis, Ranuculus) e.g. Anemone, Aquilegia Asparagales p.p. 18 Papaveraceae Onagraceae/ e.g. Iris, Maianthemum e.g. Corydalis Onagraceae Onagraceae 4 Lamiales/ 2 e.g. Epilobium 3 18 Lamiales 1 4 Fabaceae p.p. (Trifolium)/ Fabaceae p.p. (Trifolium) Campanulaceae/ Campanulaceae Asteraceae p.p. 17 phylogenetic conservatism and trait correlations with change in abundanceAsteraceae p.p. (Senecio, Solidago)/ e.g. Solidago 17 Asteraceae p.p. (Senecio, Solidago) Trait correlation 5 Apiales/ 5 Fabaceae p.p. Phylogenetic Apiaceae e.g. Desmodium (b) 2 Concord, USA Results and Discussion Our results (Fig. 1 and Table 1) indicate that change in abundance and f lowering-time response traits were phylogenetically conserved, which indicates that species evolutionary history is important to understanding community response to Table 1. Statistical tests of 1 • strong phylogenetic signalare climate change. Species that are declining in abundance more closely related than by chance. Similarly, forexpected which families of plants species that exhibit similar f lowering-time responses to temtracked or could not track perature are more closely related than expected by chance. In warmer springs contrast, latitudinal range increasingly was not phylogenetically conserved Campanulacae conservatism 16 e.g. Campanula, Lobelia 15 15 Flowering time tracking of seasonal temperature Apiaceae Shift in flowering time 1850–1900 e.g. Daucus Shift in flowering time 1900–2006 Shift in flowering time 1850–2006 Mean latitudinal range Lamiales 14 Change in abundance 1900–2006 e.g. Mentha, Verbascum n 175 319 303 271 14 414 429 Model 1 Observed rank 19 2 2,120 340 3,705 1 13 ** *** — † 12 *** Consequences of global warming? do all species have ability to track climate change to their species niche? Ranunculaceae p.p. n 175 319 303 271 414 — Model 2 6 Estimate 7 !0.48 !0.02 0.04 0.04 9 !0.10 — 6 Fabaceae p.p. n Estimate e.g. Trifolium, Vicia 7 Rosaceae p.p. * 166 !0.62 * e.g. Prunus, Spiraea *** 8 311 !0.01 * 8 Rosaceae p.p. *** 296 0.03 *** e.g. Rosa *** 253 0.03 *** 9 Sapindales, *** 362 !0.08 *** Malvales — e.g. Hibiscus, — Toxicodendron — — * *** trackers 16 Trait PhyloEcoBiogeography: Climate Change Model 3 non-trackers Chinnor n 140 140 140 140 140 — honeysuckle • invasive species were more responsive to tracking and even shifting their flowering phenology Concord Estimate difference between !1.00community *** average *0.03 * * significant *** * marginally significant 0.02 *** — — !0.09 *** — — 10 Tests used a phylogeny with branch lengths adjusted for time. The significance of phylogenetic conservatism was tested by comparing the rank of the observed 11 standard deviation (SD) of descendent trait meansRubiaceae to a null13 model based on 9,999 random iterations of trait distributions across the composite phylogeny. The 10 Cornaceae observed rank is compared with a 2-tail test significance, a P value of 0.05. Trait correlations were tested by using the e.g.of Galium, Cephalntus i.e., an observed rank of 250 11 equals e.g. Cornus comparative methods of generalized estimating equations (GEE). Estimates12describe the Primulaceae direction and magnitude of the correlation (e.g., a negative estimate Ericaceae e.g. Lysimachia #!0.1$ of mean latitude with change in abundance suggests that species from more southerly latitudes are increasing in abundance). Model 1 (univariate model), e.g. Rhododendron, Vaccinium correlation of change in abundance with each trait; Model 2 (multivariate model), correlation of change in abundance with each trait and habitat, abundance 2. Composite status phylogenies of 167Model and 323 flowering plant species fromof (a) Chinnor, UK and with (b) Concord, USA. Red (ca. 1900), flowering season, Figure and native/introduced as covariates; 3 (multivariate model), correlation change in abundance all traits and habitat, abundance (ca. 1900), flowering season, and native/introduced status as covariates (shift inresponsive flowering-time response 1850 –2006 was on excluded due to to track and blue dots at nodes represent less and more phenologically clades, respectively, based their ability its high correlation with the their other flowering flowering-time †, Ptemperature % 0.1; *, P % 0.05; 0.01; ***, P % 0.001; n % sample size.clades that have demonstrated a timesshift to traits). seasonal variation. Solid-coloured dots indicate those **, P % significant tracking response. A null distribution of mean flowering time tracking for each node in the phylogeny was generated by 9999 randomizations (‘node.mn’ in PHYLOCOM). Observed clade values that ranked less than or equal to 250 (p ! 0.05) Willis et al. indicated significantly strong trackers while values that ranked greater than or equal to 9750 (p ! 0.05) indicated significantly weak trackers. Open dots indicate marginally significant tracking clades (p ! 0.10). Each of the clades exhibiting these tracking responses is further highlighted in pink and blue and referenced numerically to its clade name. Subclades of interest are labelled with the more familiar, and more inclusive, clade to which they belong followed by pro parte (p.p.). See the electronic supplementary material, figure S1 for full taxon labels. Bar graphs (c) depict phylogenetically corrected mean differences between tracking and non-tracking clades between Chinnor and Concord. These results are normalized against the community 17030 ! www.pnas.org"cgi"doi"10.1073"pnas.0806446105 4