21 Climate Change and Forest Herbs of Temperate Deciduous Forests

Home , Anthyllis montana, Aruncus dioicus, Trillium discolor, Trillium luteum

OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN

460.1 21 Climate Change 460.2 and Forest Herbs of

460.3 Temperate Deciduous

460.4 Forests

460.5 Jesse Bellemare and David A. Moeller

460.6 Climate change is projected to be one of the top threats to biodiversity in coming 460.7 decades (Thomas et al. 2004; Parmesan 2006). In the Temperate Deciduous Forest 460.8 (TDF) biome, mounting climate change is expected to become an increasing and 460.9 long-term threat to many forest plant species (Honnay et al. 2002; Skov and Svenning 460.10 2004; Van der Veken et al. 2007a), on par with major current threats to forest plant bio- 460.11 diversity, such as high rates of deer herbivory, intensive forestry, habitat fragmentation, 460.12 and land use change (chapters 4, 14, 15, and 16, this volume). At the broadest scale, 460.13 changing climate regimes are predicted to cause major shifts in the geographic distri- 460.14 bution of the climate envelopes currently occupied by forest plants, with many spe- 460.15 cies’ ranges projected to shift northward or to higher elevations to track these changes 460.16 (Iverson and Prasad 1998; Schwartz et al. 2006; Morin et al. 2008; McKenney et al. 460.17 2011). In parallel, these climate-driven range dynamics are likely to include population 460.18 declines or regional extinctions for many plant species, particularly in more south- 460.19 erly areas and along species’ warm-margin distribution limits (Iverson and Prasad 460.20 1998; Hampe and Petit 2005; Schwartz et al. 2006; Svenning and Skov 2006; Morin 460.21 et al. 2008). 460.22 Among the plant species characteristic of TDF, forest herbs may be especially vul- 460.23 nerable to climate change for several reasons. First, many forest herbs have biological 460.24 and ecological traits that may limit the rate at which they are capable of migrating in 460.25 response to changing climate (e.g., species with seed dispersal mechanisms adapted 460.26 primarily to local movement rather than long-distance dispersal; Van der Veken et al. 460.27 2007a). Second, the fragmentation and limited connectivity of forest areas due to agri- 460.28 culture, roads, and development in the modern landscape may exacerbate the innate

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461 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:44 PM ca- nization for these species (Honnay et al. et al. nization for these species (Honnay getation to extend across large portions OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP eld-based experimentation (e.g., Van der Veken Veken der eld-based experimentation Van (e.g., cantly warmer and colder than at present (Graham 2011). For exam- For 2011). (Graham present at than colder and warmer cantly elds are providing important new insights into the relationship between between relationship the into insights new important providing are elds DEEP TIME PERSPECTIVES PERSPECTIVES TIME DEEP 1

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d d n i . 1 2 c _ elds will be key to understanding the challenges posed by modern climate change change climate modern by posed challenges the understanding to key be will elds 6 5 During much of the upper Cretaceous and Tertiary, relatively warm and wet Although the rate and eventual magnitude of modern climate change are projected projected are change ofclimate magnitudemodern eventual and rate the Although 6 7 3 8 T E M P E C R H A A T N E G E D : E C I D U O U S F O R E S T S A N D C L I M A T E ated with conservatism; temperate climatic Ricklefsforest Northern habitats conditions,and LathamHemisphere, for 1992;millionscombined Wenallowed 1999;ofwith years TDF-likePattersongreater since veandconnectivity(i.e., Givnish phylogenetic 2002). among nichelandmasses in the for modern trace TDF yearstheir plant characteristicoriginsago; communities, Manchester Ranunculaceae, and tionrise including forest1999;ofto angiospermsprominenceand Willis Liliaceae,Aceraceae, herb and toand lineages, McElwain theFagaceae,includeemerged upper many2002; such relatively Cretaceousand genera Juglandaceae,Wang as et earlythat the(~ al. 100–65 have Aristolochiaceae,in 2009). theapparently million evolutionarySimilarly, been Berberidaceae,closely diversifi associ- history lions in theof temperate have that conditions yearsclimatic spanning periods, Cretaceous upper Northernand Tertiary the into (Davis forest b eHemisphere, e n1983; bplant ple, o t hDonoghue s many ilineages g ninextricably i fi of and the haveSmith angiosperm histories linked2004; Grahamforestto extendingclimate 2011).tree changelineages Almostback 10s overall that of majormil- providemillions the of structuralyears foundation consider options for species conservation. conservation. species for options consider The plant lineages that comprise the modern TDF biome have a deep and dynamic (Delcourt 2002; Svenning 2003; Van der Veken et al. 2007a; Petit et al. 2008; Willis et al. al. et Willis 2008; al. et Petit 2007a; al. et Veken der Van 2003; Svenning 2002; (Delcourt range large-scale long-term, the about known is what review we chapter, this In 2010). biogeo- new a present and change climate past to response in herbs offorest dynamics graphic analysis investigating how contemporary distribution and diversity patterns also We dynamics. climate past these to relate herbsof may subset a forest among rare discuss how forest herb species may be affected by contemporary climate change and and developing effective conservation strategies vulnerable for effective species. plant developing and to differ qualitatively from climate dynamics in the recent geologic past (e.g., glacial cycles of the late Quaternaryimportant Period), insights into the nature of threats to impacted severely be to typesthe likely of most to species and biodiversity plant forest biogeographicperspectives and historical from drawn be may change climate rapid by s t o o d temperate , forest plants and climate change, including paleoecological (e.g., Williams s e v e ret al. a 2004) land phylogeographic research (e.g., fi Gonzales et al. 2008), et al. 2007a), comparative and der Veken studies bioclimatic Van (e.g., modeling approaches (e.g., and Skov 2004), as Svenning well as fi A synthetic view insights these vari- combining from 2011). et al. Warren 2007b; et al. fi ous challenges ofchallenges dispersal and colo long-distance herbs forest some ofdistributions geographic the Finally, volume). this 4 chapter 2002; may still be impacted by past climate change (e.g., marginalization to southern areas mak- 2007a), et al. Veken der Van 2004; and Svenning glaciations; Skov Pleistocene by magnitude the Although likely. less change climate modern to response rapid their ing of the threat to forest herb biodiversity posed by climate change is not yet fully under- 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 461 461.46 461.43 461.44 461.45 461.40 461.41 461.42 461.37 461.38 461.39 461.34 461.35 461.36 461.31 461.32 461.33 461.29 461.30 461.27 461.28 461.24 461.25 461.26 461.21 461.22 461.23 461.18 461.19 461.20 461.15 461.16 461.17 461.12 461.13 461.14 461.9 461.10 461.11 461.6 461.7 461.8 461.3 461.4 461.5 461.1 461.2 462.2 462.1 462.46 462.45 462.44 462.43 462.42 462.41 462.40 462.39 462.38 462.37 462.36 462.35 462.34 462.33 462.32 462.31 462.30 462.29 462.28 462.27 462.26 462.25 462.24 462.23 462.22 462.21 462.20 462.19 462.18 462.17 462.16 462.15 462.14 462.13 462.12 462.11 462.10 462.9 462.8 462.7 462.6 462.5 462.4 462.3 ooxfordhb-9780199837656_c21.indd 462 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 462 8 3 7 6 5 6 _ impacts of anthropogenic climate change on plant diversity and distribution in the e h t n i n o i t u . . b ) ) i 3 8 r 0 0 t 0 0 s 2 2 - i t d . s g l a n d a e i n n a t n r e e y o v t t S i a i s c t r ; i e e 3 P r v 9 e i 9 ; m d 1 4 A . 0 t ) 0 h s n 9 2 al.2006; t f al. Willis et Schwartzal.2007). ongoing 2005;For et example, et the Thuiller a 9 e r l 9 . - l o p 1 l n N k 2004; al. a et e Thomas c 2003; (Svenning n climate s changing n by i threatened n o equally t are f cies i R r e e t e e l n t g d s k o s n n a c c a a a m i e h o R e c It is clear h from climatepast episodes of m v n T d a i e i n h s t ; a t n a 2 s a e m 0 t n L t i 0 s a x l 2 e i e c ; r Q 3 t o c 9 f r f ; i 9 o u 3 n 1 o 9 r e c k 9 a g l c y 1 l o e a e i p D l l m s o ( t i r f n s d h e e u n t l m H a n k h o a c t i ; i i y b 3 R f w t 8 o i F 9 d e D F 1 s n n T D t a e T c g s a m o i n p a r v a m e h a e i t t D p e a ( o h L r long-lasting u for a potential the ; c E underscore i patterns 3 i s biogeographical 9 deep-time h These A g 9 p n 1 a i n r t r y g s e e o a l p r t o t when n t seen n u diversity o H species r c ( e contemporary in t s a differences n e striking o the r i underlie g appear to t Europe in a losses s species i ’ Quaternary c n early and a o Pliocene severe l i the Notably, g g e l r a t e h t y buffered b been have to appear losses species where Asia, eastern this of to TDF persist the in America day North eastern and Europe in extirpated lineages plant the of many from species contrast, In 2001). Manchester and Tiffney 1999; Manchester , e g n a h c s e e t i a d m u i t l s c . l ) t a 4 s c 0 - a i 0 i p g 2 r o e o l . p t o l c a e e g o s n e t n i l e o r a p u p s s d e m e r a s s i e e n l - h i i l t t c i n e W s e , p t m ) s ; f s u 9 1 i r c 9 9 h t a o 9 - 9 s n e d 1 l 1 y a o l l n p , p . a e 0 W s l n 0 e c a i 5 t ; h i d , s 2 t m u 1 t e 9 a t 2 e r 9 d n n i o 1 n o y t f f e a ~ d l m o c y a a i l y ; m e h t i s n M a r g t l n r e a G h o n a a e a i m L t a L i r m ( r a h r n P t i e L c d f o n r s r m n o i a ; p a t u a ; t t 3 e n m a 3 a s 8 s d s e s i 8 r b 9 n e c e f x 9 u n g - 1 o s r e e a 1 s i i n i u r M l m e t c s k y t s n d u o i c l x n r i t a r b f v i a e o a n v h a i a R i n a e t w , r D c o D m f h s , t ( w a i ( u o t e . s r o l t c u i g i e d e d G u o o t d a . d h d n g n l d s n u m e t n i n a o e t t i ( a a w a e v v s o s x r d h g h e d t a s a e c n t a y L b t l m c i e n r a d r h l e n e a n e c n o t e l o t f h t i h e i f i h a o a f t t r g t w c i m i t o o c e s i T d s p l c r a t l r t n e m o i e l f c e n r a c i ; w t i g t e o e 9 a h f t , f d o 9 m s a s n s ) g i e 9 i e a i o n p l 1 l i s g i a d e a c c n a r r P n l d e o u a a n e f p C . d s c e f d o W s ) r s g o i 2 a - s y n v t 0 e e t ; n i o r e s 0 y g n 9 o r r a s d 2 r a 9 i p u n n o n a l 9 t d r o t o l p 1 c e i e r a t e m v l u r F a h g o r a l o t D u t n r e h i c n T Q i f t m l o h s e c e , t e f D d 6 h ) i h o r . W t o e c o ; 2 g g n c 3 – . n s a n a e 9 3 ) i n a M r 9 . 2 o ( r 1 s 5 0 i n 0 n t n o s 2 ~ e u t e i f ( l b r v t n e i e i a i l h r s r i a k c t e d c w c o s d - l a i p i e E l R E d t c c g M a i e c m l t n i d i a c e h n l t r c p a c n a o a e i r s n l d g i i P n o l t a e l n e g i o h W c , t a ; e r n 1 v d i 0 i n 0 s u g 2 t n i r y e , t r t s d s e e h r c o n f a M l a e r o b by occupied now areas latitude high many including Eurasia, and America North of plant genera (Davis 1983; Latham and Ricklefs 1993; Svenning 2003). Fewer for- r o f r e w e F . - ) c 3 n 0 i 0 t 2 - x n e e g t n l x i a e n n n o e f i v o g S e t r ; e 3 s , e 9 n ) h 9 o 9 t 1 9 9 e 1 h o s t t f g e n d l d e n k W e a c k d i n n R e i a n ﬁ l e d n e c n n o h o a e s t i e k l m b c m P a a o h J r e e t f v h e a a t e L d h s e n t ; i t y n 3 u r e 8 b e a m including 9 t n u genera, 1 ; a r c plant 7 m e o woody 9 s i t d eight 9 i l a least 1 Dendropanax . v c u e at ) a Q r of t 5 D a r extirpation t 0 ( e e y period regional 0 s i this n l the 2 but n r a see n r America, o d r did North . e a i e eastern l B e for t n d a ( documented c e n are n e g t a h extinctions i h e plant c est t t o r t x s p e n e i E d n a v l i l t a e o p n D n c a s e including l e n c Europe, o e p o in o t s i t lineages w t s plant Carya t t e TDF a i f u f i e characteristic b i c l of h y a P numbers ; s l l large 6 e of e g 0 v tion l h 0 i a t 2 e t i v a t f n i l i o o s e n s r s i r e e h l e t g c h e u y P t o c h d t l n l a a A i r c . e ) y 3 e 0 M 0 - z 2 e n í Svenning 1993; t Ricklefs and Latham 1983; r (Davis taxa plant TDF a numerous of tion M ; 9 8 9 1 b b e W WHICH FOREST HERBS MAY BE MOST VULNERABLE E L B A R E N L U V T S O M E B Y A M S B R E H T S E R O F CHANGE? H CLIMATE TO C I H W c 2 1 - a p r i t x e l a n o i g e r r o n o i t c n i t x e e h t o t d e k n i l n e e b o s l a s a h e g n a h c e t a m i l c t s a P . i n d d , ,

Hamamelis 4 6 2 , , Platycarya , , Liriodendron in situ in , , Pterocarya evolution of species climatic tolerances (Huntley and d n a y e l t n u H ( s e c n a r e l o t c i t a m i l c s e i c e p s f o n o i t u l o v e , , Magnolia , and d n a , change and future projections that not all spe- suggesting long-term niche conservatism in conservatism niche long-term suggesting Sciadopitys , , Tsuga (~ 2.6 million years ago to present), the , and upward of 80 other woody woody other 80 of upward and , (Latham and Ricklefs 1993; ; 3 9 9 1 s f e l k c i R d n a m a h t a L ( nal gla- 110/23/2013 5:19:45 PM 0 / 2 3 / 2 0 1 3

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463 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:45 PM cally, sub- cally, cant new threat threat new cant ions of small-ranged species may soon greater greater risk of extinction due simply to develop between the locations of many many of locations the between develop species traces to a number of ecological onset of new a climatic qualitatively regime the locations of climatically similar areas in rized by lower abundances and smaller popu- me species this is likely to result in population OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP usting usting their distributions in response to anthropo- change (Latham and Ricklefs 1993; Svenning 2003). 2003). Svenning 1993; Ricklefs and (Latham change 3 6 4

d d n i . 1 2 c _ 6 5 Of greatest concern in the face of modern climate change are species with limited 6 7 3 8 W H Y RANGES? M I G H T S M A L L - R A N G E D S P E C I E S H A V E S M A L L of endemism, the most commonly cited are species’ innate biological or ecological characteristics (e.g., competitive inferiority or association with uncommon habitats; evolutionary recent their 2004), Lavergne 1989; Baskin and Baskin 1978; Daubenmire origin (Stebbins and 1965; Major Levin 2000; or Lesica et 2006), endemism al. due to the contraction of a formerly more extensive range (Daubenmire 1978). These three Ecologists Ecologists have long recognized that the restricted endemic distributions plant of species small-ranged may be the Wherry 1944; Stebbins and Among potential Major 1965; drivers Daubenmire outcome 1978). of a variety of causes (Willis 1922; Clark 1998; Williams et al. 2004). Nevertheless, the substantial numbers of forest plant plant offorest numbers substantial the Nevertheless, 2004). al. et Williams 1998; Clark extirpations and extinctions to the linked during the late Tertiary and early Quaternary suggest that not climate abrupt to all resilient equally forest plants are poleward poleward range shifts of and mammal, many bird, insect taxa suggest that some relatively vagile species are already adj al. et Zuckerberg 2005; al. et Hickling 2003; Yohe and (Parmesan change climate genic 2009; Breed et Although 2012). al. similar range shifts in response to modern climate record paleoecological the plants, forest for documented well been yet not have change suggests that some species may be capable of rapid relatively range adjustments (e.g., Schwartz Schwartz et Without al. successful 2006). long-distance dispersal to track shifting climate zones as they move poleward, populat be exposed to novel climatic regimes that fall outside the range tions they of exist under currently; for climatic so condi- 2004). (Thomasdeclines or extinction et al. stantial geographic disjunctions are likely to small-ranged current species’ ranges and pres- between disjunctions Such 2006). al. et Schwartz 2004; (Thomas future the al. et ent and future habitat areas are less likely for widespread where species, at least some portions of these broadly distributed rangesspecies’ are likely to remain climatically suitable into the buffering future, against climate-driven threats (Thomas et al. 2004; et et al. 2006). This characteristic, combined with the geographic clustering of populations, may expose small-ranged species to stochastic population or processes to chance regional events introduc- (e.g., drought, tion of novel pathogens; Gaston 2003). In addition to risk factors that may be inher- modern small poses change to climate range a ently signiﬁlinked size, to many small-ranged, endemic species (Thomas et al. 2004, 2011). Speciﬁ of extinction projected for small-ranged and biogeographical factors. For example, macroecological studies have frequently detected a positive correlation between range size and local abundance, small-ranged such that lation sizes than widespread species (Gaston 2003), a result that has been apparent in several plant-focused studies (Thompson et al. species 1998; Murphy et al. 2006; Pocock are often characte geographic distributions, such as endemics and other small-ranged species (Thomas et al. 2004; Parmesan 2006; Schwartz et al. 2006; Thomas 2011). The increased risk 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 463 463.46 463.43 463.44 463.45 463.40 463.41 463.42 463.38 463.39 463.36 463.37 463.33 463.34 463.35 463.30 463.31 463.32 463.27 463.28 463.29 463.24 463.25 463.26 463.21 463.22 463.23 463.18 463.19 463.20 463.15 463.16 463.17 463.12 463.13 463.14 463.9 463.10 463.11 463.6 463.7 463.8 463.3 463.4 463.5 463.1 463.2 464.2 464.1 464.46 464.45 464.44 464.43 464.42 464.41 464.40 464.39 464.38 464.37 464.36 464.35 464.34 464.33 464.32 464.31 464.30 464.29 464.28 464.27 464.26 464.25 464.24 464.23 464.22 464.21 464.20 464.19 464.18 464.17 464.16 464.15 464.14 464.13 464.12 464.11 464.10 464.9 464.8 464.7 464.6 464.5 464.4 464.3 ooxfordhb-9780199837656_c21.indd 464 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 464 8 3 7 6 5 6 _ the of comprehensiveTo reviews no America? North beenhave eastern date,there in climate from change, risk what do we know increasedabout the current small-ranged distributions of forest herbs at be to expected are species small-ranged that Given ally linked in forest (Van herbs der Veken et al. 2007a). al. et 2004), factors,(Thomas limitation tocaus- be dispersal sizeand like range small risk change climate key for potential the highlighted have studies These vertebrates). or wind via (e.g., dispersal longer-distance for adaptations exhibiting seeds with cies adaptations for dispersal had signifi morphological lacking those and ants) via (e.g., dispersal local to adapted seeds with herbs forest European that found (2007a) al. Vekenet der Vanexample, For 2010). some forest herbs (e.g., Skov and Svenning 2004; Van der Veken et al. 2007a; Bellemare scales that dispersal geographic limitation may also contribute to limited sizerange in larger at studies from increasing is 1994; evidence volume),Matlack this forests; 16, chapter secondary of recolonization post-agricultural (e.g., timeframes rela- over short c scales tively local e on p focused have s herbs forest a in limitation dispersal seed s ing a investigat- studies most l Although volume). l this 2007a;16, al. e chapter Vekenet der w s a , (e.g.,manyMatlack al.1994;Bellemareforest et herbs 2002; Verheyen s al.et 2003; Van signifi c to lead i may agents t dispersal s suitable of i the absence and r dispersal, seed e long-distance for t adaptations c morphological of a lack a r tion, a h c s e graphic history. For example, i studies have shown that factors such as c low seed produc- e p s havewill climate modern to limited change. track ability tor infl Jump 2011). In general, the potential for dispersal limitation to be a key historical fac- dispersal limitation (Svenning and Skov 2007a; Vanto linked der Vekenbe could et al. ameliorated 2007a; Hampe have and conditions after ranges paleoendemics’ of sion distributions (Daubenmire 1978; Levin 2000), it is also evident that the limited expan- restricted species’ these to leading decline and fragmentation range of dynamics the on focus frequently paleoendemics of considerations Although 2008). 2005; al. Kooymanet Rosetto and (Rossetto paleoendemics some of distributions restricted the in involved factor key a as suggested been also has limitation Dispersal 2006). al. et (Lesica ranges their expand and disperse to time havelimited may simply had species range diffi and expansion dispersal seed inter-site making likely habitat, unsuitable of matrix a in scattered widely often are patches habitat bedrock),suitable serpentine (e.g., habitats unusual to linked are distributions whose endemics ecological of case the In 2008). al. et Van2007a; RossettoSkovVeken der 2007a; and al.Svenning et 2005;Kooyman manydistributions of geographic restricted small-ranged plants (Kropf et al. the 2002; Rossetto and to contributing factor a be also may limitation dispersal seed that ity possibil- the raised increasingly have studies species, plant endemic of ranges small additio In 2001). Cruzan and Estill 1978; “paleoendemics,”demics,” respective and endemics,”“ecologicaltermed “neoen- been have species endemic of classes general WHERE ARE SMALL-RANGED FOREST HERBS IN N I S B R E H T S E R O F D E G N A R - L L A M S E R AMERICA? A NORTH E EASTERN R E H W c 2 1 Dispersal limitation range of size for endemic forest herbs could be traced to innate . i n d d uencing the small range size of many endemic plants suggests that these species

4 6 4 cult. For neoendemics, evidence suggests that some recently evolved recently some that suggests evidence neoendemics, For cult. cantly smaller geographic ranges than ranges cantly related smaller geographic spe- ts of regional landscape structure and biogeo- and structure landscape regional of ts y Sebn ad ao 16; Daubenmire 1965; Major and (Stebbins ly n to these traditional explanations for the for explanations traditional these to n cantdispersal limitation for 110/23/2013 5:19:45 PM 0 / 2 3 / 2 0 1 3

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465 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:45 PM ed as endan- as ed (Flora of North cally on the distribution of ies. ies. Further, conservation options for Flora Flora of North America ecies ecies were also included if their habitat n in this survey, we visually inspected all ded was also described as occasionally occur- cally on small-ranged plants associated with with associated plants small-ranged on cally situated withinsituated matrix a deciduous forest (e.g., OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP cally on the TDF biome. Prior studies by Stein et al. (2000) cant issue in quantifying species distributions as almost all forest 5 6 4

d d n i . 1 2 c _ 6 5 ned “small-ranged” plant species as those with distributions including 70 or fewer “small-ranged” ned For For each small-ranged herbaceous species with a distribution centered in eastern In In the analysis presented here, we have focused specifi 6 7 3 8 ring outside forest habitats in meadows, open rocky areas, wetlands, or along road- distribution at a BONAP provided data for Of plant species in Canada are sides. note, did this however, level); (county U.S. the within than level) province (i.e., scale coarser not become a signifi herbs of distributed south with substantially ranges small were border. Canadian the 2003; Weakley 2011). 2003; Species Weakley were selected for inclusion if their habitat descriptions included deciduous forest or woodland, or mixed deciduous-coniferous forest (e.g., hardwood-hemlock or oak-pine forest). Sp specialized but stillwas more typically shaded ledges, woodland clearings, forest edges, forested seeps and stream A subset banks). of the forest herb species inclu population biology and threats to federally listed forest herbs). herbs). forest federally listed biology to population threats and North America, we reviewed habitat information to identify those that were associated with deciduous forest habitats using the America editorial committee 1993+) and et key al. 1968; Gleason regional and Cronquist 1991; 1999; Yatskievych references Wunderlin and Hansen (e.g., Radford d e fi Although counties. many plant species U.S. with small ranges are classifi gered or threatened at the federal or state level, our species selection process did not impor- an as size range consider we rather, criterion; a as status listed current consider current of species’ regardless ofchange, face climate the ofrisk in future correlate tant for a review of this volume, 4, chapter Harris and legal Pimm also status 2008; see (cf. To identify To appropriate species for inclusio plant species distribution maps developed by the Biota of North America Program Kartesz 2010) for (BONAP; species with geographic ranges in centered eastern North con- a on updated are and ) www.bonap.org ( online available are maps These America. pres- the in distribution used the maps available; become records new as basis tinuing ent analysis were accessed from BONAP in 2010. For the purposes of this we survey, small-ranged forest plants may include some approaches (e.g., assisted colonization) that may be less feasible for species more associated unusual withand spatially other, limited habitats where endemics are often such found, as serpentine barrens or lime- glades. stone small-ranged forest herbs associated with TDF habitats in eastern North America. Tennessee), in Tennessee), addition to a limited number of hotspots in TDF (e.g., the southern Estill Mountains; Appalachian and AlthoughCruzan 2001). these earlier studies have been key to mapping the distribution and diversity of plant endemics in general, biome-centered a survey focusing specifi TDF has not been conducted. Such a study will be crucial in the context of climate change, as the unique ecology and biogeographic history of forest plants may dispose pre- them to climate-related vulnerabilit distribution of small-ranged forest plants (i.e., endemics) or analyses of patterns of endemism focused specifi and Estill and Cruzan (2001) have surveyed patterns of eastern North endemism America, but neither focused in on forest portionshabitats in detail. These ofinves- tigationsscrub and sand hill vegetation in central Florida, open cedar glade habitats in central highlighted numerous “hotspots” of endemism in non-forest habitats (e.g., 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 465 465.45 465.46 465.47 465.48 465.42 465.43 465.44 465.39 465.40 465.41 465.36 465.37 465.38 465.33 465.34 465.35 465.30 465.31 465.32 465.27 465.28 465.29 465.24 465.25 465.26 465.21 465.22 465.23 465.18 465.19 465.20 465.15 465.16 465.17 465.12 465.13 465.14 465.9 465.10 465.11 465.6 465.7 465.8 465.3 465.4 465.5 465.1 465.2 466.9 466.8 466.7 466.6 466.5 466.4 466.3 466.2 466.1 466.34 466.33 466.32 466.31 466.30 466.29 466.28 466.27 466.26 466.25 466.24 466.23 466.22 466.21 466.20 466.19 466.18 466.17 466.16 466.15 466.14 466.13 466.12 466.11 466.10 ooxfordhb-9780199837656_c21.indd 466 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 466 8 3 7 6 5 6 _ km drawn almost entirely from m o r f y l e r i t n e ﬁ the to t species s o m l a species,respectively. Pteridophytes ly and n w a r d were two latter the of representatives Notably, percent). 11 ~ spp., (20 Cyperaceae and percent), 11 ~ spp., (21 Melanthiaceaepercent), 11 ~ spp., (21 Lamiaceae per-cent), 18 ~ spp., (34 Asteraceae included: species the of percent 50 ~ for accounting These species represent taxa from 38 families 21.1).and 87 genera,(appendix with four of America these families North eastern in forests deciduous to native species herb these quantiles. box),and 75thquantiles(outer edgesof and dashed “whisker” databeyond of therange linesmark 2010. Box plot (top) size depicts mean (diamond), range size medianline), (vertical range the 25th North Biota of ProgramAmerica (BONAP) mapsaccessed county-level speciesdistribution in sizes were estimated U.S. asthetotal area of count size was ~ 280,000 km 280,000 ~ was size bedrock dolomite on habitats glade and a median range size of ~ 61,448 km 61,448 ~ of size range median a county-level distributions. Range sizes exhibited a positively skewed distribution, with centroid additionally, a species; that by each calculatedwas species as summation the by occupied area range total The (GIS). system information geographic a into tized digi- were maps distribution county-level BONAP the herbs,forest small-ranged 189 eastern North areAmerica relatively widespread. appalachianum herbs like e k i l L. areas (estimated range 2.3, ~ 2.7, km million 3.5 and s b r e h forest large-ranged of ranges the than smaller magnitude of order an almost still are as such analysis, our in included species small-ranged widespread most the of sizes range the even Notably, Pennsylvania. southwestern to Carolina North western from mid-Appalachians the in woods mountain to native species a Britton, FIGURE c 2 1 I ttl te rtra ulnd bv rsle i a e o 19 ml-agd forest small-ranged 189 of set a in resulted above outlined criteria the total, In o rvd qatttv etmts f ag sz ad egahc oiin o the for position geographic and size range of estimates quantitative provide To . 2 i n for r o f d d

21.1

4 0 Onosmodium decipiens Onosmodium 6 6 forest species included in the this 189 study. small-ranged herb Range sizes of Range Podophyllum peltatum Podophyllum in habitats forest lycophytes with and associated ferns most Pryer),as nalanalysis ( 50000 2 for the relatively more widespread widespread more relatively the for Trillium Botrychium mormo Botrychium J. woodlandnative Allison,open narrowtoendemic a Range Size(km L., 150000 Asarum canadense Asarum and 2 (ﬁ was estimated for each range based on these on based range each for estimated was g.21.1). Minimum range size was1,600~ cophytescontributed twoonly small-ranged in Bibb County, Alabama. Maximum range range MaximumCounty, Alabama.Bibb in Carex ies occupiedies by each species, determined as from of the areas of all the counties the areasoccupied of of 2 ) , with 21 and 18 small-ranged forest small-ranged 18 and 21 with , W. H. Wagner and and WagnerW.H. 2 L., or r o , . L , respectively). 250000 Meehania cordata Meehania Sanguinaria canadensis Sanguinaria Gymnocarpium Gymnocarpium 20 40 M. cordata M.

Count (Nutt.) 110/23/2013 5:19:45 PM 0 / 2 3 , , /

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467 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:46 PM cial 0 1 sp 2–3 spp 4–5 spp 6–10 spp 11–20 spp 21–30 spp 31–40 spp 41–50 spp 51–59 spp Last Glacial Maximum Temp Decid Forest Small-Ranged Forest Herb Spp. Richness species per county). The results of this of this results The county). per species

g. 21.2). At the broadest scale, small-ranged for- small-ranged scale, broadest the At 21.2). g. rn in portion counties some and of biome, TDF the OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP gs. 21.2 and 21.3). Although these northern areas areas northern these Although 21.3). and 21.2 gs. communities, almost all of the species found north north found species the of all almost communities, 7 6 4

d Distribution richness and of herb North small-ranged 189 eastern in species forest d n i . 1 21.2 2

c _ 6 5 To assess overall patterns of small-ranged forest herb distribution and diversity in eastern eastern in diversity and distribution herb forest small-ranged of patterns overall assess To 6 7 3 8

FIGURE and the Last Glacial Maximum (LGM; blue line). County-level richness of County-level small-ranged line). blue forest (LGM; Maximum Glacial Last the and of recorded species low a herbs to high a zero ranges of Carolina from North western in species 59 of much formerlyacross the glaciated northe Boundaries of Embayment. TDF Mississippi and Plain Coastal the along U.S. southeastern the surfi state-level boundary from derived was LGM the (1999); al. et Ricketts follow biome geology maps. America relative to the distribution of the Temperate Deciduous Forest biome (TDF; biome line) green Forest Deciduous distribution the to of Temperate the America relative est herbs are relatively common in the southeastern U.S. and lower Midwest, but are almost almost are but Midwest, lower and U.S. southeastern the in common relatively are herbs est entirely upper absent Midwest, often from include ofTDF and the areas adjacent estLGMwell-developed northhaveherb relativelyCanadaofspecies the forestLast(fi that large Glacial herbwere geographic the Maximum focus (LGM)rangesof this in when theanalysis. comparedNortheast, to the small-ranged for- North eastern across herbs forest America,small-ranged of small-rangeddiversity hotspots and pronounced distribution the both that with show analysis patterning, biogeographical rangemarked exhibit America North species maps for and richness coldspotsthe 189 (no. species of ofendemic small-ranged were species compiled richness in a GIS (fi to create a map of 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 467 467.9 467.6 467.7 467.8 467.2 467.3 467.4 467.5 467.1 467.21 467.18 467.19 467.20 467.15 467.16 467.17 467.12 467.13 467.14 467.10 467.11 468.36 468.35 468.34 468.33 468.32 468.31 468.30 468.29 468.28 468.27 468.26 468.25 468.24 468.23 468.22 468.21 468.20 468.19 468.18 468.17 468.16 468.15 468.14 468.13 468.12 468.11 468.10 468.9 468.8 468.7 468.6 468.5 468.4 468.3 468.2 468.1 ooxfordhb-9780199837656_c21.indd 468 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 468 8 3 7 6 5 6 _ hotspots can be somewhat subjective when confronted with the complex diversity pat- t a p y t i s r e v i d x e l p m o c e - h v t e s h t e i d w u . l s d c r e n e t i v n i o t r r s f e g n w n o d o c i l M a n e s r h f e w f w o u e l l v b i l d t r a n c o m a e s j s . b e S u f p . s o o U l t s n a s r h t fi g e w o ( n t e p i s m s n c a o t o a e s o i f h h - g t e h e , u b r t M o e small-rangedforest r diversity G of distinctivenessrelativelygeographic s v hightheir and to L n o e i a n h t e c t e c h s h n t a t s i f t t o , terns h evident o ourin results, s f e wefocus herethree on c prominent p i h o areas that m stand outdue u s d c o s t u h i o , m b t , y . h a u . l M d s o fi S F l G a s s ( . D L a n o U T c a r s m i C e c n e k h e h a i r p h t o c e 0 a t t t e 0 r n s f p 2 g i f a o s , o o e endemism 1 e of M levelsmoderate b to h lowheterogeneous h – more g backgroundof pattern G a r t as s t 0 L e u n 0 r h o o l 8 o e s i a n h t t r g t s e r e n s e h o i a r f t p y e o o l r f n n a h i r d , t e e e n r n g m i o i n fi o n a a i l e ; r b p m m M - k o G l F l L i l D a 0 b a T t e 0 m s d 9 F s fi a e i – D h o s 0 T f g c t n 0 o n i 8 i e f s n h m d o n i t k n o a e i s m 0 t f t n e 0 x o tobeassociated a i r withpatches 1 TDF-like of e habitat r g incooler andmore mesic sites onthe t t e r < e n n h a m e s e t m o c t d i n ; outsideorthesouthern x on i of theTDFbiome;boundary n of these outlying b species o tend ) e o r c y r e F t r l h h D n a a t g T e d i u i c n small-ranged t o h e species ( u (~ a s 8 percent) h o included p e in y this t b analysis s e v have r range centroids a h located a M f h t e d G o g L n r g u s a n e o n l o d b o l i i a s M e t n G h r L i t o p e m e h k t , t i t 6 but p no c small-ranged e 8 species distributions s a l approached 1 e f this d limit. In contrast, d d some d areas n a e I r d t e t a g s c r o t e l a m regions h glaciated e d formerly t i in herbs o forest s r endemic e of t richness g n low n e of pattern a c This r ( e M v G a L h e h t f o northern Minnesota, Wisconsin, and Michigan) has a range situated substantially north distance:centroidskm(meanhaveLGM range438situatedthe substantially of south the TDF biome in the southeastern U.S., of boundaries the LGM. ~ 800–1,200 km south of forest species are 16 small-ranged the centroids herb found near or beyond the southern of the LGM of (i.e., than 186 km north located further –186 km on the LGM; of however, Canada ~ 800–900 km north areas of species centroids no small-ranged are axis extends the LGM, to –800 km, of or 800 km north as the TDF biome extends northward into The meandistance from centroids range to theLGM north was438km(±224SD). Thedistance the LGM,of regions; glaciated the LGM formerly is set to within 0 on the distance axis. boundary the LGM,of outsideregions; glaciated formerly species in eastern North America. Positive values indicate centroids range that are situated south boundary, only y l n o , y r a fi d n ; u D o S b 4 2 2 ± FIGURE 80–500 –800 c 2 1 - h t r o n t s o m m o r f s b r e h t s e r o f d e g n a r - l l a m s f o e c n e s b a l a r e n e g e h t o t t s a r t n o c n I speciesAmong189theincludedanalysis,this in almostspp.,percent)(183all97 ~ . ot South North i n d d

21.3

4 6 8 Range centroid distances to the LGM forest for 189 small-ranged herb boundary g.21.3). Of the six species with range centroids falling north of the LGM orcim mormo Botrychium Range CentroidDistancetoLGM(km) 200204060801000 800 600 400 200 0 –200 g. 21.2). Although the criteria for deﬁ LGM ( peioht ntv t sgr al frss in forests maple sugar to native pteridophyte (a negative indicate values centroids located north -agd oet eb iest, s well diversity, as herb forest l-ranged x axisinthisﬁ g. 21.2). Indeed, 16 of the ningand delineating gure). In contrast, ve species g. 21.3). 10 20 30 40 110/23/2013 5:19:47 PM 0 / 2 3

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469 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:47 PM To r r . L . C . Magnolia g. 21.2). 21.2). g. A r n o t t . g. 21.2). 21.2). g. ora, such as as such ora, Liatris gholsonii Shortia galacifolia g. 21.2). In total, 119 of the the of 119 total, In 21.2). g. Torreya taxifolia (Vail) Woodson, occurs in this region in the Florida panhandle panhandle Florida the in region N a c z i a n d of Arkansas and Missouri (ﬁ among these 119 species, 18 have ranges holds for small-ranged forest herbs, with ecies with overlapping ecies distributionswith in overlapping both Michx. is found only in cool, mesic forests at clude counties in and around the Apalachicola Apalachicola the around and in counties clude in eastern North America (peaking at 59 species 59 at (peaking America North eastern in OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP Carex Carex thornei ApalachicolaRiver Matelea alabamensis Nuttall ex Chapman, and , t h e Interior Highlands oridana Diphylleia cymosa Diphylleia Taxus ﬂ 9 6 4

d Southern Appalachians Southern d n i . 1 2 c _ We a t h e r b y , 6 5 6 7 3 8 The Interior Highlands Hotspot Hotspot Highlands Interior The The Interior Plateau ablyHighlands in Arkansas,less hotspot,attention Missouri, including in theand partsbotanicalextreme of easternthe and Ouachita ecological Oklahoma, Mountains literaturehas receivedand Ozark on consider- forest plant diversity to the Apalachicola River hotspot ( est herb River species area. the Apalachicola haveImportantly geographic distributions southern rangesthough,River forest Appalachian areathat that thisherbs inrepresentsalso hotspot in includeMountains, this is a comprised area,southernmostcounties or only the primarilyfarther twoadjacent extension (7 to percent)of thePiedmont. species northor disjunctare infor Of narrowcentral which thestation 29endemics Alabama, small-ranged in restrictedthe entirely The Apalachicola River Hotspot Hotspot River The Apalachicola The Apalachicola Alabama small-ranged and Gadsden Riversouthwestern forest areaCounty, ofherbs, GeorgiaFlorida, the peaking Floridais and the atDecatur 21region panhandle sp County, with and Georgia.the adjacentnext Overall, highest southeastern 29 richness small-ranged of for- (Weakley core 2011). area western to south Virginia, western and Virginia West from TheMountains, ofAppalachian southern to highNorthspatial the diversityCarolina, extentsouthern aandof broaderedges thisendemism of hotspotzone the inof Appalachian alsohighthe southerndiversityseems Plateau remarkable: Appalachian extends in northeastern along Beyond Mountains most the of Alabama theof mid- (fi 189 small-ranged overlap that the are species hotspot).Southern entirely high (63 ForAppalachian elevationsrestrictedpercent) & example,A. Gray reviewed hotspot;in tois thethisa well-knownsouthern inregion this (i.e., Appalachiansurvey narrow 15 have percentendemic Mountains,distributions ofnative the while speciesto that just occurring six counties in the in the region ter of Estill plant counties anddiversity Cruzan extreme in and of 2001). westernnorthern small-rangedendemism with This NorthoverlappingGeorgia intrend foresteastern Carolina, andclearly distributionsherbs westernNorth eastern anywhere AmericaSouth Tennessee, in Carolina (e.g.,western Stein including southwesternNorth etCarolina; al. the 2000; Virginia,highest fi richness and h e r b s : t h e Hotspot The Southern Appalachian Previous studies have highlighted the southern Appalachian Mountains as a major cen- Anderson); one additional species, and adjacent Georgia, and the area, as hotspot well a as sdoes h ein i onealso county include in severaleastern narrowGeorgia. endemics Notably inthough, its woody the Apalachicolafl River 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 469 469.41 469.39 469.40 469.38 469.35 469.36 469.37 469.32 469.33 469.34 469.29 469.30 469.31 469.26 469.27 469.28 469.23 469.24 469.25 469.22 469.19 469.20 469.21 469.16 469.17 469.18 469.13 469.14 469.15 469.10 469.11 469.12 469.7 469.8 469.9 469.4 469.5 469.6 469.3 469.1 469.2 470.2 470.1 470.43 470.42 470.41 470.40 470.39 470.38 470.37 470.36 470.35 470.34 470.33 470.32 470.31 470.30 470.29 470.28 470.27 470.26 470.25 470.24 470.23 470.22 470.21 470.20 470.19 470.18 470.17 470.16 470.15 470.14 470.13 470.12 470.11 470.10 470.9 470.8 470.7 470.6 470.5 470.4 470.3 ooxfordhb-9780199837656_c21.indd 470 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 470 8 3 7 6 5 6 _ (Pittman and Bates 1989), and d n a , ) 9 8 9 1 s e t a B d n a n a m t t i P ( Berkeley County, South Carolina, also stand out as areas with relatively high num- m u n h g i h y l e v i t a l e r h t i w s a e r a s a t u o d n a . t s s b r o e s h l a t s , e a r n o i f l o d r e a g C n a h r t - u l o l S a , m y s t n f u o o C s r y e e b l e k r e B and Carolina, North County, Pender counties, plain coastal two Finally, Pennell). Kral & V , ) 1 9 9 1 s e t a B d n a l a r K ( s e t a B . M V. & l a r K range size and latitude (fi latitude and size range signifi highly a contrast, In 0.05). > , . g . e ( y k c u t n e K n r e h t r o n t n e c a j d a and Illinois, have Indiana, Ohio, River Valley southern in Ohio species the along centered herb ranges forest small-ranged of number a north, the to Further Duncan. T s a h c u s s e i c e p s b r e h t s e r o f d e b i r c s e d y l t n e c e r s a l l e w s a , r o l y Ta Delphinium newtonianum range size with increasing latitude, often referred to as Rapoport’s Rule (Lomolino o n i l o m o L ( e l u R s ’ t r o p o p a R s a o t d e r r e f e r n e t f o , e d u t i t a l g n i s a e r c n i h t i w tern (e.g., increases in species niche breadths e with latitude; Stevens 1989), the z relatively i s et al. 2006). Although a number hypotheses of have e been advanced to explain this pat- g n a r o commonly the with consistent is lation l the a in c east, River i the Savannah upper the h to along counties p Further Carolina South a several northeast. region, r Piedmont the g to o hotspot e Appalachian g Southern the e from distinct is e that endemics r local and regional h for hotspot a as t emerges Alabama and e U.S. h southeastern t the central-western in County Tuscaloosa of around area an these, Among Midwest. lower parts other o in t apparent also are overlap- 10–15 distributions) n (e.g., ping diversity o of i peaks lower t with i hotspots secondary d of d number a n I Secondary Hotspots to the west in our study area might infl might area study our in west the to availability water and rainfall decreased that expectations despite apparent, was tude - d longi- to relative u size range in trend l no species, 189 the c boundary.Among LGM the n i , s b historical factors: centroidrange longitude, centroid latitude, and centroid r distance to and e geographical three and h size range between correlations analyzed t also we s herbs, e r o f d forest small-ranged diversityof and distribution the in overalltopatterns addition In e g n a r - l l a m s f Trends inRange o s e i t i s n e d h g i h y l e v i t a l e r t i b i h x e o s l a d like e endemics, h narrow some s ing r e t a w cies (25 percent) associated with the Interior Highlands hotspot are narrow endemics s c i m e d n e w o r r a n h e t h r e t o r N a f o s n t a r o t e p h n t s c e s t u n a o s o e h p s m s n e o d i i c n c a e y r l p t s e h s s i i g g s s . r i e e t r n a H d n i e o l u h v i l , r c i g c t y o i d e n o l i r r t i p l r b o s a e r b p e t i t e e r s s o t n h h e t e h n I t h o h a h r T t t e e s s t h . n v e b s y t n i i r u e e o R o r s k h i f o t a g g f a a i l e n f w o r i o d c e r s e s i s d r a g i h i s e u n c h t r t c s a a t o p a c , i r l p m i o t s t - a s o c - s y s l p c t o o r l u A l o c s i a j a h s s F e n m a c h a o s s i r . t t e ) m o a s i t e j f n e d i c n d a o a r e h e e n m h a t t p c e t c k s r i t r a e ) n r n w e e p s i o a t h p 8 a r t s t 2 s 5 r e r o t r n 2 r a o u e a ( n p o e w h k m h o t r s i , t s l i A e d d w i n d , n m y c y o n a o l c t a t , r t e n y s f h S u t t g o c i t i d C . n s i l e a i r y s t regional c t fl e a r , i s v r l e i r i a e o m d e d v s o m e y i g A a w b t b o n y s r h o d l s e M e l e h ; z a l n t i c e i s r h t a e t s s e t e e e c n r i a e o o c r h n f e a t p h s c d o e 9 t g 1 h n ( g r a u e r o h - h t l t r l l u a A f m s s a e f r o a n a h t c 2 1 . i n d d

4 7 0 ora than in the other two hotspot regions: Seven the 28 of small-ranged spe-

Size D. M. Moore, and d n a , e r o o M . M . D g. 21.4b; F 21.4b; g. Stachys iltisii Stachys rlim discolor Trillium Oxalis illinoiensis Oxalis cantpositive correlation wasapparent between 1 ,187 uence range size for forest herbs (ﬁ 11.5, = y itntv htpt dsrbd bv, a above, described hotspots distinctive ly Polymnia cossatotensis Polymnia bserved biogeographic trend of increasing of trend biogeographic bserved J. Nelson (Nelson 2008). . ) 8 0 0 2 n o s l e N ( n o s l e N . J Solidago ouachitensis it is geographically and physiographi- p = 0.0009, R 0.0009, = Wray ex Hook. and d n a . k o o H x e y a r W Schwegm., , . m g e w h c S Carex latebracteata Pittman & V s e t a B . M V. & n a m t t i P Hydrophyllum brownei Hydrophyllum 2 = 0.06). This corre- This 0.06). = C. E. S. T . J . R & r o l y Ta . S . E . C Penstemon deamii Penstemon . persistens T. g. 21.4a, 21.4a,g. Waterfall, , l l a f r e t a W 110/23/2013 5:19:48 PM 0 / 2 p 3 /

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471 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:48 PM g. 21.4c), 21.4c), g. = 0.1 for for 0.1 = ,187 1 = 0.07; ﬁ 0.07; = 2 indicated in each panel in each indicated 2 = 0.0002, R 0.0002, = = 0.0 p 2 0 –500 = 0.06 = 0.07 ) -values and R R and -values p = 0.73 R 2 2 ° ) p ° p = 0.0002 R p = 0.0009 R t more closely by a regression in range = 14.0, 14.0, = . The . 2 500 uous decline tracking latitude north of the 1,187 Latitude ( Longitude ( OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP 1000 Centroid Dist. to LGM (km) 70 80 90 100 25 35 45 to ~ 280,000 km to 2

9 8 7 6 9 8 7 6 9 8 7 6 1500

13 11 10 14 13 12 11 10 12 14 14 13 11 10 12

LN (Range Size) (Range LN LN (Range Size) (Range LN LN (Range Size) (Range LN (c) (a) (b)

1 7 4

d Correlations between natural log-transformed range size and species’ range centroid range centroid natural between log-transformedCorrelations range and species’ size d n i . 1 21.4 2

c _ 6 5 6 7 3 8 FIGURE panel A; 11.5 for B; 14.0 for C. C. for 14.0 B; for 11.5 A; panel LGM, suggests an important historical component to the pattern in our study area (cf. (cf. area study our in pattern the to component historical important an suggests LGM, Cowling and Samways 1994; Dynesius and Jansson 2000; Jansson 2003). Consistent the trend within range this size possibility, is ﬁ boundary(F LGM the to distance centroid abrupt truncation in the distribution and richness of small-ranged species near the rather than a boundary, more contin LGM an analysis that the into account takes irregular lobes border and ofmajor southward range size varied from ~ 1,600 km km 1,600 ~ range varied size from longitude (panel A), latitude (B), and distance to the LGM boundary (C) for 189 small-ranged LGM the to and distance (B), latitude A), longitude (panel herbs study, this in included small-ranged the forest Among America. herbs North eastern in forest are derived from simple linear regression; the associated F statistics are as follows: F as follows: F statistics are the associated simple linear regression; from derived are 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 471 471.12 471.13 471.14 471.9 471.10 471.11 471.7 471.8 471.4 471.5 471.6 471.1 471.2 471.3 472.2 472.1 472.45 472.44 472.43 472.42 472.41 472.40 472.39 472.38 472.37 472.36 472.35 472.34 472.33 472.32 472.31 472.30 472.29 472.28 472.27 472.26 472.25 472.24 472.23 472.22 472.21 472.20 472.19 472.18 472.17 472.16 472.15 472.14 472.13 472.12 472.11 472.10 472.9 472.8 472.7 472.6 472.5 472.4 472.3 ooxfordhb-9780199837656_c21.indd 472 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 472 8 3 7 6 5 6 _ havesubstantially herbs forest small-ranged few very that apparent is scale,it graphic on post-glacial migration rates and so-called refugia.cryptic First, at the broadest geo- 21.3).and 21.2 maypattern This several interesting suggestivebe processes of bearing (fi LGM the of north areas most from herbs forest small-ranged of absence the to U.S.southeastern contrastedthe when lowerparticularly Midwestand surprising, are diversityapparent acrossmuch of forestherb small-ranged low-to-moderateof levels above. the Indeed, described hotspots of number limited the exclusivelyto restricted Jackson et al. 2000), but exhibits low forest small-ranged (fi herbs diversity of al.1998; et Cain 1975; Delcourt and Delcourt (e.g., species forest temperate for gium fr has region Valley:RiverThis LowerMississippi the in seen is hotspots diversity herb forest small-ranged and refugia glacial et al. 2001). One notable exception to this pattern correspondenceof between putative Near1998;Robison and Carlton(e.g., region the nativeto species animal fromdence 1999), al. et (Rickettsalthough most recent refugiumresearch has focused Pleistocene-era on biogeographic and phylogeographic a evi- as described been also has study identifi hotspot Highlands Interior The 2008). al. et Gonzales 2005; al. et (McLachlanLGM the during persistedhave may species plant forest temperate some dence points to the southern Appalachian Mountains as an area where populations of evi- genetic population Similarly,increasing2001). Cruzan and Estill 1949; (Thorne example, the Apalachicola River area has long been hypothesized as refugiuma glacial For America. North eastern in refugia Pleistocene-era important as suggested ously Skov and 2007a). (Svenning refugia glacial former these around or in regions to restricted haveto appear remainedspecies forestplant of subset a of al. ranges et 2004),the but out these of southern areas (Davis 1983; Prentice et al. 1991; Cain et al. 1998; Williams distributions Holocenethe eras,early and Pleistocene late the in 2001; climate Cruzan of amelioration and the With (Estill 2007a). Skov LGM and Svenning the during habitat suitable of areas small to south contracted species zone temperate of ranges the when havedeveloped to thought hotspots are Such 2009). Diadema and Médail 2001; Cruzan and (Estill refugia glacial vey,frequentlyhas Pleistocene-era beentaken indirectas locationsthefor of evidence sur- this of results the in seen LGM,as the of south the to far endemism of hotspots concentratio the Similarly, 2011). al. et Sandel 2007; al. et Finnie 2003; Jansson 2000; Jansson and Dynesius 1994; Samways and (Cowling species endemic of distributions the on effect major a havehad ciation other areas of the Northern in Hemisphere and endemism suggests that past of climate change patterns and gla- on observations with consistent is America North eastern of portions glaciated formerly most from herbs forest small-ranged of absence The have species these ex actually that distributions of few very while even LGM, the toward increase to tend herbs forest small-ranged (fi advance glacial last the POSED BY MODERN CLIMATE T A M I : L S C S E N C R O E R D P O M O T Y N B R E D T E T S A O P P M O R F c 2 1 It is also clear from the results of this study that not all small-ranged forest herbs are identifi hotspots major the of three All . i n d d

4 7 2 g. 21.2). Overall, these results show that the range sizes of of sizes range the that show results these Overall, 21.2). g. o mn salrne seis n distinct in species small-ranged many of n

ed in this study correspond to areas previ- areas to correspond study this in ed

CHANGE of many temperate plant species expanded species plant temperatemany of INSIGHTS INTO THREATS equently been mentioned as a likely refu- likely a as mentioned been equently tend north of this boundary. of tend north edinthis g. 21.2). 110/23/2013 5:19:48 PM 0 gs. / 2 3 / 2 0 1 3

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Hexastylis spp., spp., Geranium Geranium macu- ed for the analy- the for ed eld for these spe- these for eld , g. 21.3). This pat- This 21.3). g. seeds dispersed by Carex Trillium eld observations (e.g., Matlack Matlack (e.g., eld observations 300 km from the LGM, well outside the the outside well LGM, the from km 300 Sanguinaria canadensis cally, 43 of the small-ranged forest herbs herbs forest small-ranged the of 43 cally, , context including wide-ranging congeners or or wide-ranging congeners including context cies with limited dispersal ability (e.g., species ability dispersal with cies (e.g., limited OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP of small-ranged species identiﬁ Asarum Asarum canadense 3 spp., various Lamiaceae and Ranunculaceae spp.). Similarly, the pres- 7 4

d d n i . 1 2 L.). Rare long-distance dispersal events clearly need to be invoked to account c Trillium _ 6 5 Prior studies have linked small range size in forest herbs to biological and ecological ecological and biological to herbs forest in size range small linked have studies Prior In particular, the dispersal and range dynamics suggested by the results ofpres- the results the by suggested dynamics range and dispersal the particular, In The The second pattern evident in our results with implications for estimating migra- 6 7 3 8 l a t u m for the distribution patterns seen among these wide-ranging species, and subsequent studies have documented potential mechanisms (e.g., have not formally reviewed the life history traits of the 189 species included in the pres- the in included species 189 the of traits history life the reviewed formally not have as ent little study, published data is available on these relatively rare, range-restricted species. it However, is striking that a large number of these forest herbs come from spe include to known genera or families with ant-dispersed seed or no obvious mechanism of dispersal: spp., (1998), the present study focused on small-ranged endemics, a group that has typically has that group a endemics, small-ranged on focused study present the (1998), is thoughit even literature, paleoecological and dispersal plant the in overlooked been among such species long-term where dispersal limitation of range size is a reasonable 2007a). et al. Veken der Van 2004; and Svenning (Skov hypothesis We 2007a). al. et Veken der (Van ability dispersal and production seed limited like traits were were common large-ranged species with distributions extending well into formerly glaciated regions (e.g., Cain by in et contrast the al. to species considered However, 2003). et al. Vellend deer; cies and the substantial distances many must have migrated during the Holocene to current reach range boundaries in the Based north. Cain on et these al. discrepancies, (1998) concluded that rare long-distance dispersal events likely enable rapid migration and range shifts in forest herbs (cf. Clark 1998), even for species that otherwise appear to be severely dispersal-limited based on though, Notably almost 1994). all of fi the forest herbs by considered Cain et (1998) al. for for other temperate forest plant species (e.g., Cain et al. 1998; Clark 1998; Williams 2004). et al. ent survey appear to diverge most (1998) strikingly al. et from Cain conclusions change. drawn climate by to response Cain in migration et herb al. forest regarding (1998) reviewed on literature the dispersal ability of 28 herbsforest and highlighted the mis- between the match limited seed dispersal distances reported in the fi expanded expanded or shifted their distributions into formerly glaciated regions in the north; only six of the 189 species (3 percent) included in this analysis had range centroids situated north of the LGM, and most species range centroids were situated substantially south of this boundary (mean distance to LGM = 438 km; fi to seems and deglaciation widespread since years 15,000 ~ nearly despite emerges tern stand in marked contrast to the relatively rapid northward range expansion inferred confamilials (cf. Lavergne et al. 2004; Van der Veken et al. 2007a). 2007a). et al. Veken der Van 2004; et al. Lavergne (cf. confamilials tion capacity of forest herbs was the close proximity of some distributions small-ranged to the species LGM boundary. Specifi (23 percent of total) had range centroids ence of only two ferns and lycophytes (i.e., taxa that typically produce large quantities quantities large typically that taxa produce (i.e., of lycophytes ferns and ence two only of wind-dispersed spores) in the set sis seems telling. In contrast, the large number of small-ranged Asteraceae (34 spp.), a family often characterized by wind-dispersed propagules, was surprising. Clearly, further research on the trait characteristics of these small-ranged species is needed, phylogenetic comparative a in especially 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 473 473.45 473.46 473.47 473.48 473.42 473.43 473.44 473.39 473.40 473.41 473.36 473.37 473.38 473.33 473.34 473.35 473.30 473.31 473.32 473.27 473.28 473.29 473.24 473.25 473.26 473.21 473.22 473.23 473.18 473.19 473.20 473.15 473.16 473.17 473.12 473.13 473.14 473.9 473.10 473.11 473.6 473.7 473.8 473.3 473.4 473.5 473.1 473.2 474.2 474.1 474.47 474.46 474.45 474.44 474.43 474.42 474.41 474.40 474.39 474.38 474.37 474.36 474.35 474.34 474.33 474.32 474.31 474.30 474.29 474.28 474.27 474.26 474.25 474.24 474.23 474.22 474.21 474.20 474.19 474.18 474.17 474.16 474.15 474.14 474.13 474.12 474.11 474.10 474.9 474.8 474.7 474.6 474.5 474.4 474.3 ooxfordhb-9780199837656_c21.indd 474 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 474 8 3 7 6 5 6 _ for detected been have Mountains Appalachian southern the and haplotypes unique between associations Similar range.species’ the in south farther seen those to relative divergent were that haplotypes in areas of Kentucky and Tennessee, as well as in the southern Appalachians, documented (2008) al. et Gonzales example,For 2008). al. et isolated in distinct glacial refugia during the wereLGM or earlier glacial maxima (Gonzales that populations of descendants the represent to believed are lineages genetic lower Mississippi River Valley (McLachlan et al. 2005; Hu et al. 2009). These divergent in types temperate forest plant populations well to the north the of Gulf Coast and the haplo- unique documented have studies phylogeographic recent LGM,as the during Mississippi River Valley; 1975, Delcourt and Delcourt lower 1987; Davis 1983). the and Coast Gulf the (e.g.,U.S. southeastern the in refugia cited glacial major traditionally as areas the outside refugia, northern cryptic in LGM the through (fi However,LGM fi this the to closer distributed species by exhibited size range larger the tially northward during the Holocene Epoch, a dynamic that would be consistent with havesubstan-species expanded or shifted of subset this of ranges geographic the that major hotspots identifi 1,000 m/yr or more to keep pace with modern climate change, but even the fastest the even but change, climate modern with pace keep to more or m/yr 1,000 approach to need will rates migration plant that haveprojected studies Indeed, some be expected to migrate in response to modern climate change (McLachlan et al. 2005). record is now being reevaluated, implications for how critical species can with rapidly - pollen the l on based species o plant forest many for ; estimated p originally potential 5 tion 0 e 0 h 2 t . Cain et y al. m 1998; Clark 1998; see also MacLachlan l et (e.g.,al. Valley 2005). l o RiverAs such, the a Mississippihigh migra- e lower r or k f Coast Gulf i t the from l e dispersal g long-distance n . , i n ) M s a 8 G s l L 9 i h 9 substantially lower m than what has previously c 1 e a been h have L e may . rates migration t c r post-glacial l that imply M a would a boundary LGM ( the fi of g i t n s c e i n b e o r r p n i u e S i t d h a a C l t s o s a t , s e e i . i r r m g cant o a implications for i estimates post-glacial of . migration rates (McLachlan et f al. d l 2005). e n ( e a t y s s l s e o n e h , y m o i t t c i e e , n s p tree c s i TDF n s u s better-documented e d e of d s o l those e r from d i p extrapolated r w c a typically o were e g y l dynamics p e e s R h , t range their and s record, pollen . the t on based e studies ) e n paleoecological to z largely invisible 1 have been s a i herbs 1 u forest l s studies, 0 a p genetic recent 2 c these to n e prior grasses, such, F trees, As o b sedges. n many D and of i a T pollen t d v wind-dispersed a r o abundant l f o more r the u c P o to compared p e as o r d e p n c n a n e e l l s l y e a t r m t p s a e B e o h t t e d u e d t c e t e d not generally have record pollen the on focused studies paleoecological that in tic” (fi r o f m forest herbs small-ranged relativelya of arearichness with u high northern a as analysis our i in emerged g also that u region a f Minnesota, southeastern e and Wisconsin r western l a i c gla- a 2005).Moreof strikingly,Provanevidence presented and genetic (2011) Beatty c 2 g. 21.2). 1 Increasing genetic evidence points to the existence refugia northern such of cryptic The new evidence for cryptic northern refugia during the LGM may have signifi have may LGM the during refugia northern cryptic for evidence new The “cryp- as to referred are data genetic these from inferred refugia northern The . i n d d cally, the persistence temperate of forest plant populations a within few 100 km

4 7 4 nding may also indicate that some small-ranged forest herbs persisted Monotropa hypopitys Monotropa ed in the southeastern U.S. (ﬁ Acer rubrum Acer and L. unglaciated the Area”L.“Driftless in south- of Fagus grandifolia Fagus ed quantities of insect-dispersed pollen, insect-dispersed of quantities ed been inferred based on models assuming g. 21.3). This pattern may suggest al. et (McLachlanEhrh. Trilliumcuneatum g. 21.4). g. Raf. . f a R 110/23/2013 5:19:49 PM 0 / 2 - 3 / 2 0 1 3

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475 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:49 PM , , native to the Apalachicola River hotspot, particular, particular, because hotspots of endemism and OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP survive via local elevational shifts rather than Torreya Torreya taxifolia (Delcourt 2002; Hampe and Petit 2005; Wilson et al. 2005; Ashcroft Ashcroft 2005; et al. Wilson 2005; (Delcourt and Hampe 2002; Petit 5 7 4

d d n warming i . 1 in the past, their ranges may now be poorly positioned to withstand future ed in this analysis may be associated with areas that exhibit some resilience to 2 c _ 6 5 In In the face of such climate-driven threats, conservationists have traditionally Interestingly, there Interestingly, is evidence that hotspots of endemism tend to occur in areas 6 7 3 8 C O N S E R V A T I O N I M P L I C A T I O N S 2007). Unfortunately, this approach may prove ineffectual for species that are severely severely for that species ineffectual are prove may this approach Unfortunately, 2007). dispersal-limited, or for those whose present ranges are separated and by large potential expanses of future unsuitable habitat habitat (Thomas begun to have some et researchers these challenges, Given this volume). 4, chapter al. 2004; Thomas 2011; shifts shifts in regional climate zones, with the potential for also see off effect; entirely of escalator so-called the tops the southern (i.e., mountains some habitats to disappear Delcourt and Delcourt 1998). stressed the importance of habitat corridors and landscape connectivity to facilitate natural dispersal and range shifts (Hunter et al. 1988; Hannah et al. 2002; Hunter forest forest herbs to areas native with limited topographic heterogeneity Gulf(e.g., Coastal moun- in those to relative risk increased at be may U.S.) midwestern ofportions Plain, tainous areas, as successful tracking of climate envelopes for the former species will likely require larger latitudinal displacement of ranges (cf. Sandel et al. 2011). At the other small-rangedextreme, species linked to high elevation habitats in the southern Appalachian Mountains may also face severe habitat loss due to upward elevational climatic stress, such as mesic sites, river valleys, and north-facing slopes, may allow for local persistence despite changing climate (Jansson 2003; Ashcroft 2010; Sandel et al. 2011). Consequently, it is possible that the hotspots and small-ranged species identiﬁ near-term climate change; however, the magnitude of modern climate change may eventually overwhelm such environmental buffering. In this context, small-ranged of one 2004). Schwartz and Martin 2004; (Barlow change in part climate be linked to may narrow endemic, that have historically permitted some resilience to climate Ashcroft 2010; Sandel et regionsFor 2011). al. example, with change substantial topographic (Jansson 2003; heterogeneity may allow species to large-scale migration; similarly, the presence of microhabitats that may moderate 2010). 2010). Consistent with this prediction, relict populations of a number of boreal and range southern their at recruitment failing or limited exhibit already species plant TDF edges in Europe (e.g., García et al. 1999; Hampe and 2002; Arroyo Mejías et al. 2002, 2007; Castro et al. 2004; Beatty et al. southern the 2008). at plants In forest small-ranged of eastern dynamics population the North on focused America, have few studies decline severe the that suggested have researchers some but biome, TDF the of margins in the southeastern U.S. would likely facilitate conservation planning under more sta- conservation planning under facilitate more likely would in the U.S. southeastern ble climatic conditions, the rapid climate change projected for coming decades may substantially complicate this goal. In diversity tend to be localized to southern areas where TDF species survived climatic cooling climatic migrations of the late Pleistocene and early Holocene now appear to have been on the the on been have to appear now Holocene early and migrationsof Pleistocene late the 2008). al. et Petit 2005; al. et oforder (McLachlan less or m/year 100 Although the co-occurrence of many small-ranged forest herbs in regional hotspots 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 475 475.43 475.44 475.45 475.40 475.41 475.42 475.37 475.38 475.39 475.34 475.35 475.36 475.31 475.32 475.33 475.28 475.29 475.30 475.25 475.26 475.27 475.22 475.23 475.24 475.19 475.20 475.21 475.16 475.17 475.18 475.13 475.14 475.15 475.10 475.11 475.12 475.7 475.8 475.9 475.4 475.5 475.6 475.3 475.1 475.2 476.2 476.1 476.45 476.44 476.43 476.42 476.41 476.40 476.39 476.38 476.37 476.36 476.35 476.34 476.33 476.32 476.31 476.30 476.29 476.28 476.27 476.26 476.25 476.24 476.23 476.22 476.21 476.20 476.19 476.18 476.17 476.16 476.15 476.14 476.13 476.12 476.11 476.10 476.9 476.8 476.7 476.6 476.5 476.4 476.3 ooxfordhb-9780199837656_c21.indd 476 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 476 8 3 7 6 5 6 _ the potential for the types of large-scale translocations envisioned to ensure long-term m r e t g - n g i n t o i l m i e l r u , s y n l e l a o t t n e d m e e n r o c i . n s ) i i 1 v 1 y n 0 l e 2 n o s s n a o r m i u o t c h a c T n c o ( o o t l s t r s e h a n m g B a i i r g m d t e n r a t e e a l c h k t a i t c i c t i b s a e r a - m n t h e i i a g l m p r c e a k a l x r l w b e i e a d l K f n t a o o i s , t i r u n . r s e s o g b r e d i . i e p n t e l f d u y a ( i o r t l - u o l u i n q a n p e d o e n v o o h n i fi i i i p t o t r species appears t v increasingly , tenuous (e.g., c Svenning a and a Skov u h 2004, r e r s 2007a,b; Schwartz t B b s u g o t p l i i t s n f n a a l d c r - a e d t b n i t s m h m l a n a a d s e c e n a e t e i t i o i g m r s d n r c t t n e n p e o i e n l i t o c p v e o e t a r o n s t H h i m y i t t e o t m i l v a p ; i l n l l h t 5 l c e i a n t e n 0 a c i h e 0 n f n i t r n 2 c i o p t r s e i y i a y u t e n . s l t h t c f w e l e e p w i t g a c k m h e o r , h i u d s b b p o t r l t s l o r f i o e s u r e e w d a o m h h y h l w u e t t u r e m i s t n o a h u r a s a w n s t y i b C e n o l r i r r s o i n e b l a o ; e i t o d i i n f 3 g t u o l u o 0 d a d l m i d q i 0 e l e o u e e t 2 u v s q g u p e e i e n l c o g t m a o i p n l e s v r n m r a o u e - i a H p r o d l n i n l , e y e r ; y a a r r d f a 7 l m a a o v 9 r , s r 9 a e a a y s s a 1 l t f d g n n i a o n o t f i o m m u l n o i g i i e o o e t r S l l b e c s a a c o n e e l m r . i o r e u ) y l e p g p 3 e l c l e n o 0 g l e a r a p 0 n a d p r 2 a p g r s i n d n t e c n e n i n g v n o a n e d i i t i r e r s r , a r y d a p t h u - G e s p c e , g u a t s n s r a s n a - g m t o r f o i i i l e l t e g c y d s o r n e i a o t h b y r a t e o , h b p y d t g e o t d d l a e o o c h o c i s h e d i i n l l i b e a k e t i c s l n e e e d h i t v e s i f i n e v E e v r e s e r p o t forest that may herbs be threatened by climate modern change. l a i t small-ranged for tool conservation potential a n as colonization assistedter, discuss we e t o p e biome.or fi region Inthe any particular of candidatespecies m o s likely history,and ecology, biogeographic s the in grounding clear lack r to tended have e f f discussions these such, as o globe; the around from drawn examples a extreme o of with range s illustrated or l nature in a hypothetical largely been ) have n colonization assisted o i of discussions t most date, However,to a 2010). Collins v and Minteer 2009; r Simberloff e s e r fi p and Ricciardi 2008; Hellmann Mueller d and o (e.g., species translocated among siveness y l O r c ; inva- for potential the 7 a regarding particularly conservationists, 0 and i ecologists among 0 v 2 , . . l g a . e t ( e s n k for potential a n e the future l adaptive reduce evolution in respo a m and h b wild o the c in s a extinction d to L e species f consign c e might o alone a s M n ( n o i i l t a e technique this a using Pritchard2009),but and (Li climate-threatenedspecies and c g rare v i a r t r e c o s a t e s r r p p m i m r e e b t - g n o may arboreta and gardens botanic l in them) within diversity genetic the (and species e h t indefi the and impossible, r be may o ranges former compromised f y r a s neces- be may conservation situ ex to unconventionalapproach This 2011). Thomas 2008;al. et Hoegh-Guldberg 2007;al. et (McLachlanfuture the in changes climate as cally, but where they are expected to histori- survive as self-sustaining,occurred naturalized populations not have they where regions to species translocating intentionally et McLachlanal. 2007; 2004;Hoegh-Guldberg et al. Martin 2008; Thomas 2011). and Assisted (Barlowcolonization proposes change climate rapid to due extinctions cies spe- avoid to relocation managed or colonization assisted for potential the consider WOULD ASSISTED COLONIZATION OF SMALL-RANGED D E G N A R - L L A M S F O N O I T A Z I N O L O C D E T S I S S FEASIBLE? BE A HERBS D FOREST L U O W c 2 1 However, the applicability of equilibrial range models to small-ranged TDF plant TDF small-ranged to models range equilibrial of However, applicability the debate vigorous sparked has colonization assisted of possibility the Nevertheless, . i n d d

4 7 6 nse to climate change (Davis et al. 2005). that species translocated beyond their range range their beyond translocated species that growthrates(Webb 1986; Woodward 1987; ag mdl wud ed o ugs that suggest to tend would models range y on species’ ranges typically assumes that assumes typically species’ ranges on y species, as reintroduction into climatically eld 2009; Thomas 2011). Long-term seed ies? Most species distribution models in models distribution species Most ies? ue to abiotic or biotic limits. Even with eld2011). Overall, these “equilibrial” such aspeciescould besuccessfully nal sections of this chap- this of sections nal nitemaintenance of 110/23/2013 5:19:49 PM 0 / 2 3 / 2 0 1 3

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477 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:49 PM u- (L.) cant Torreya Torreya W a l t e r , Aesculus Aesculus (L.) Salisb., xed limits deter- ect ﬁ ect (Steud.) Sleumer, Catalpa bignonioides Hyacinthoides Hyacinthoides non-scripta Eranthis Eranthis hyemalis (L.) Pers. germinated and success- (L.) Pers. Lam., cally, for cally, plant species with signiﬁ L.; Lid and Lid 1994; Stace 1997; Skov time since amelioration of past climatic stress, stress, of climatic amelioration past since time le over substantially greater spatial scales than OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP Leucothoe Leucothoe fontanesiana (Walter) Fernald, Jeffersonia diphylla ve-year period in forest habitats 200 km beyond the species’ Aristolochia Aristolochia macrophylla Rhododendron Rhododendron ponticum Aruncus Aruncus dioicus L., (Ker. Gawl.) Torr., Gawl.) (Ker. Torr., L., 7 7 4

d d n i . 1 2 c _ 6 5 Even Even more strikingly, numerous incidences of small-ranged forest plant species Other sources of information on plant species’ climatic tolerances and the potential potential the and tolerances climatic species’ plant on informationof sources Other 6 7 3 8 and Svenning 2004). Although such patterns have not been as extensively documented documented extensively as been not have patterns such Although 2004). Svenning and for forest plants in notable eastern America, cases North of small-ranged forest herbs and woody species naturalizing in areas far to the north of their natural ranges have been observed (e.g., Dicentra eximia range limits have been documented (Gleason and Cronquist 1991; Skov and Svenning Svenning and Skov 1991; Cronquist and (Gleason documented been have limits range 2004; Kartesz 2010). In Europe, a number of plant species endemic to areas around Pleistocene-era glacial refugia in southern observed to readily and naturalize in TDF forests south-central of northwestern Europe (e.g., Europe have been hippocastanum Lilium martagon in in AlthoughEurope. horticultural observations do not provide reliable information on the role that biotic factors (e.g., competitors, pollinators, pathogens, herbivores) theydistributions the limiting of in wild, the in might species small-rangedplay plant range-restricted species. per se is not limiting for many do demonstrate that climate escaping from horticulture and naturalizing in forest communities well beyond their plant species’ distributions are routinely tested et der (Van Veken al. 2008; Sax et al. 2013). In particular, the horticultural trade includes numerous small-ranged natu- of their north more or forest km 1,000 to 100s many grown commonly are that species ral ranges in eastern North America (Dirr 1998; Cullina 2000, 2002; Sax et al. 2013). grown, were plants native that found (2008) al. et Veken der Van reviewby a Similarly, on average, ~ 1,000 km north of their natural range edges in the horticultural trade actually occupied (cf. Skov and Svenning 2004). assisted Consequently, colonization efforts for such species might be feasib equilibrial range standard models. by be predicted would unplanned or accidental many the effortsare colonization ofgeographic scale assisted evident “experiments” in horticulture, where the climatic limits on numerous native natural range edge in the northeastern U.S. Van Similarly, der et Veken al. (2007b) data on an extra-rangepresented transplant experiment almost ear- initiated 50 years lier that showed long-term survival and expansion of edge range natural its beyond km 100 ~ to up areas in populations Rothm. ex Chouard These empirical studies suggest of that the extent Europe. potentially in northwestern suitable habitat for many dispersal-limited forest herbs may greatly exceed the area of geographic ranges is not widely acknowledged by paleoecologists (e.g., Webb 1986; 1986; of geographic rangesWebb is not widely paleoecologists (e.g., by acknowledged Prentice et al. 1991; Williams et al. 2001; but see Davis 1986), empirical evidence for plant TDF among increasing is 1986) Davis (sensu type“disequilibrium” this of range der Van 2004; Skov and Svenning 2004; Svenning and Skov 1980; Holland (e.g., species et al. 2007b; Bellemare Veken 2010). For example, Bellemare (2010) found that seeds of the herbant-dispersed forest fully established over a fi et al. 2006; Van der Veken et et al. der 2007a). Veken al. Specifi 2006; Van dispersal limitation, current range boundaries might not refl mined by environmental factors, but rather slow-moving colonization fronts infl rates, dispersal species’ by largely enced and the geographic locations of former refugia (Holt et al. 2005; Svenning and Skov 2007a,b; Bellemare 2010). Although the potential for long-term dispersal limitation 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 477 477.45 477.46 477.47 477.48 477.42 477.43 477.44 477.39 477.40 477.41 477.36 477.37 477.38 477.33 477.34 477.35 477.30 477.31 477.32 477.27 477.28 477.29 477.24 477.25 477.26 477.21 477.22 477.23 477.18 477.19 477.20 477.15 477.16 477.17 477.12 477.13 477.14 477.9 477.10 477.11 477.6 477.7 477.8 477.3 477.4 477.5 477.1 477.2 478.2 478.1 478.45 478.44 478.43 478.42 478.41 478.40 478.39 478.38 478.37 478.36 478.35 478.34 478.33 478.32 478.31 478.30 478.29 478.28 478.27 478.26 478.25 478.24 478.23 478.22 478.21 478.20 478.19 478.18 478.17 478.16 478.15 478.14 478.13 478.12 478.11 478.10 478.9 478.8 478.7 478.6 478.5 478.4 478.3 ooxfordhb-9780199837656_c21.indd 478 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 478 8 3 7 6 5 6 _ forsomespecies. andeffectivenecessary strategy conservation among TDF small-ranged plants and, as a result, assisted colonization could be both a suggest that large-scale dispersal limitation may be a relatively common phenomenon evidence of lines various These 2010). Kartesz 2004; Martin and Barlow 1997; Case approach to test hypotheses about demographic history in forest herbs. . s b r e h t s e r o f n i y r o t s i h c i h p a r g o m e d t u o herbs. b Most notably, and within seed-sowing experimental include might suchefforts a s e forest in assumption this test directly toresearch s experimental more for need great a e h t o p is al.2006),there y et Schwartz 2005; Thuiller and Guisan al.1995; et (Huntleyclimate h t s e current t with equilibrium distributional presuming models on based are change mate o t h c a o r p p a this used yet have that studies any of aware not areWe history. demographic ering large datasets on nuclear DNA would be a substantially more powerful tool for uncov- diversity(e.g., Grifﬁ allozyme on phylogeographicfocused studies genetic population and cpDNAhaplotypes using studies of focus the been have plants forest some Although populations. tories should leave distinct signatures in DNAsamples of sequences drawn from these his- demographic contrasting margins,these range southern at declined margins,but years; of reviewed in Moeller 100s–1,000s et al. 2011). (e.g., If scales time longer substantially over ranges geographic across dynamics population of evidence examine to opportunity an provide studies genetic options. considered as candidates for management, translocation, or other ex situ conservation to species TDF small-ranged multiple on ously simultane- run be potentially could they time-consuming, albeit conduct, to simple technically be would studies these likely.Because be may declines such that suggest studies other and this in detected patterns biogeographic the though even America, (i.e., rates growth population ing or species small-ranged populations, herb forest of studies demographic any of aware not Weare endemics). TDF woody ous numer- as well as 21.1, appendix in listed species (e.g., species plant of numbers tial substan- on needed is research descriptive basic decline, climate-driven of clear evidence demonstrating species to limited be should translocation and intervention at line against which future population dynamics base- a establish to order in species plant TDF small-ranged of populations existing forest herbs, climate rapid change, and conservation. Here we outline what we see as some 2013).of the al. et Sax 2010; Collins and Minteer 2007; al. et (McLachlan implementation for considered be should they before investigation substantial require will nization, unconventionalresponses to challenges, conservation newthese such assistedas colo- biodiversity.Likewise, herb forest to change climate modern by posed threat of tude magni- the bettertounderstand needed research considerablestill that is evident Itis a i l o f i x a t O P E N Q U E S T I O N S A N D R E S E A R C H O P P O R T U N I T I E S S E I T I N U T R O P P O H C R A E S E R D N A S N O I T S E U Q N E P O c 2 1 Third, given that most analyses projecting plant species’ responses to future cli- future to responses species’ plant projecting analyses most that given Third, ﬁ to contrast in Second, monitor and document to needed is effort research major a foremost, and First . i n d d

4 , , 7 8 Trillium luteum Trillium n and Barrett 2004; MacLachlan et al. 2005; Gonzales et al. 2008), (Muhl.) Harbison; Gleason and Cronquist 1991; Case and Case 1991; Cronquist and Gleason Harbison; (Muhl.) edbsd eorpi suis mlclr population molecular studies, demographic eld-based λ < 1) at southern range margins in eastern North eastern in margins range southern at 1) <

populations have expanded at northern range otherwise, that have demonstrated declin- demonstrated have that otherwise, key open questions relating to small-ranged could be gauged. Because any attempts determine which, if any, should be 110/23/2013 5:19:50 PM 0 / 2 3 / 2 0 1 3

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479 Climate Change and Forest Herbs 0 2 / 3 2 / 0 110/23/2013 5:19:50 PM past by uenced for this type of investigation in light of modern climate change will likely expe- likely will change climate modern on invaders of intercontinental origin (e.g., of intercontinental invaders on ting species’ migration potentials and evalu- and potentials migration species’ ting OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN 2013, 23 Oct Wed – FIRSTPROOFS, PROOF UNCORRECTED OUP bored within and among populations in species’ species’ in populations among and within bored c diversity in the future (Hampe and Petit 2005; th and Watson 2006; Van der Veken et al. 2007b; Bellemare 2010). 2007b; 2010). Bellemare et al. Veken der Van 2006; th Watson and 9 owering in many species), even if migrating populations were to per- 7 4

d d n i . cant need for further insight to warm-margin the southern, nature of species’ 1 2 c _ 6 5 Finally, it has become clear that historical post-glacial range expansion has involved involved has expansion range post-glacial historical that clear become has it Finally, Fourth, Fourth, whether forest herbs migrate naturally in response to climate change or 6 7 3 8 S U M M A R Y Research increasingly indicates that dispersal limitation may be a major factor control- factor major a be may limitation dispersal that indicates increasingly Research geographiccurrent the the ling that and species distribution plant of forest numerous distributions of range-restricted still species be many may strongly infl native native ranges and how different genotypes may perform in novel northern environ- experiments; garden common through (e.g., variation genetic such Identifying ments. et 2011) al. may be Fournier-Level cf. key to designing conservation successful efforts and preserving valuable intra-specifi 2007). et al. McLachlan rience natural selection on ecologically important traits (Geber and Dawson 1993; Etterson and Shaw 2001; Davis et al. 2005). For example, northward migration will involve substantial shifts in photoperiod (an important cue for development, dor- mancy, and fl fectly track a particular set of climatic factors. It is important, har currently then, is variation genetic what to understand such such biotic interactions within and beyond natural species’ range limits. Insight into predic to key be will dynamics biotic these ating risks with colonization. associated assisted evolutionary not change, simply migration (Davis and Shaw 2001; Davis et al. 2005), to response in migrating populations that and and Hellman and 2008; Hellman Simberloff This might difference et 2012). al. be due to a range of key a is herbivores) pathogens, (e.g., enemies natural from escape example, for factors, factor that has been linked to invasiveness among intercontinental exotics (Mitchell and 2003; Power Carpenter and Cappuccino 2005), but this ecological phenomenon An important focus for be the with less may typeslikely intracontinental movements. of forest herb seed-sowing experiments described above will be documentation of communities communities will be colonized by new species in coming decades. Such intracontinental movements have received relatively little attention in the invasion biology lit- primarily focused been has which erature, chapter this It 12, is volume). not yet clear if intra-Mack et versus 2000; intercon- al. intracon- that evidence some but indicates comparable, directly are invasions tinental tinental movement of plants does not commonly lead to invasive behavior (Mueller distribution limits. If, as predicted by some warm-margin ecological range edges theory are determined primarily by (MacArthur biotic factors (e.g., competi- 1972), moder- to response limited be actually may there climate, than rather herbivory), tion, ate levels of climate or change, species responses could be confounded or accelerated 2010). al. et Putten der (Van interactions biotic complex by plant forest many colonization, assisted via intentionally moved are species threatened beyond beyond current range boundaries to assess plant performance and its relationship to and Geber 2005; Schemske and Angert 2004; al. et Eckhart (e.g., factors environmental Eckhart 2005; Griffi target clear a are edges range northern Although also is there efforts, colonization assisted or migration offuture direction probable the a signifi 9 9 1 0 8 7 9 - b h d r o f x ooxfordhb-9780199837656_c21.indd 479 479.46 479.44 479.45 479.42 479.43 479.39 479.40 479.41 479.36 479.37 479.38 479.33 479.34 479.35 479.30 479.31 479.32 479.27 479.28 479.29 479.24 479.25 479.26 479.21 479.22 479.23 479.18 479.19 479.20 479.15 479.16 479.17 479.12 479.13 479.14 479.9 479.10 479.11 479.6 479.7 479.8 479.3 479.4 479.5 479.1 479.2 480.2 480.1 480.28 480.27 480.26 480.25 480.24 480.23 480.22 480.21 480.20 480.19 480.18 480.17 480.16 480.15 480.14 480.13 480.12 480.11 480.10 480.9 480.8 480.7 480.6 480.5 480.4 480.3 ooxfordhb-9780199837656_c21.indd 480 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9

9 Community Dynamics and the Role of Disturbance 480 8 3 7 6 5 6 _ Nomenclature follows Kartesz (2010). Range statistics were derived from county-level l e v e l - y t n u o c m o r f d e v i r e d e r e w s c i t s i t a t s e g n a R . ) 0 1 0 2 and Case 1997; Yatskievych ( 1999; Wunderlin and Hansen 2003; Weakley 2011). z s e Case 1991;Cronquist t and Gleason al.1968; et Radford(e.g.,sources regional various r a K s w the from o drawn l was information l Habitat www.bonap.org). o see f (BONAP; Program America e r u t a l distribution maps c developed for each species by Kartesz n (2010) and the Biota of North e m o N America. North eastern in Forest TemperateDeciduous with associated herbs forest small-ranged 189 for habitat and longitude, and latitude centroid range area, Range Analysis Lab. this projectof was provided by Lilly Dalton and Jon Caris in the Smith College Spatial components GIS the with ValuableKartesz.assistance John and (BONAP) Program America North of Biota the of efforts the through compiled data distribution detailed plant the without possible been have not would here presented analysis graphic biogeo- Verheyen.The Kris Hermy, and Vellend, Martin Mark Somers, Paul Peter Marks, Svenning, Jens-Christian Agrawal, Anurag Geber, Monica including uscript, valuab provided colleagues Numerous forest species.vulnerable herb particularly some for considered be to need may colonization, assisted like options, signifi a be to fi these Given 2004). al. et (Thomas risk this signifi may limitation dispersal large-scale long-term, of challenge to be at increased from risk climatemodern change due to smallsize, range the added priori a predicted be would species endemic these of many Holocene.As the during many endemic species have exhibited relatively that limited migration and suggesting possibility,range expansion this with consistent highly are herbs forest small-ranged of survey our from emerging patterns biogeographic change.climateThe of episodes APPENDIX I D N E P P A S T N E M G D E L W O N K C A c 2 1 . i n d d

4 8 0 Flora of North America North of Flora cant threat to forest herb biodiversity, forestherb threatto cant unconventionaland conservation

21.1 21.1

for species covered by published volumes and from and volumes published bycovered species for le comments on earlier versions of this man- this of versions earlier on comments le ndings,modern climate change is likely cantly compound cantly

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5 Species Family Range Area (km 2 ) Range Centroid Range Centroid Habitat Description 6 _ c 2

1 Latitude (º) Longitude (º) . i n d d

Aconitum reclinatum A. Gray Ranunculaceae 43681 37.736 80.554 Rich cove forests, seeps & shaded ravines, mtn 4 8 1 woods Aconitum uncinatum L. Ranunculaceae 167349 37.011 81.422 Mesic woods, seeps & clearings Actaea podocarpa DC Ranunculaceae 107768 37.649 81.176 Moist, rich wooded slopes & coves Actaea rubifolia (Kearney) Ranunculaceae 38063 36.564 85.945 Rich cove forests over calcareous bedrock Kartesz Ageratina luciae-brauniae Asteraceae 11840 36.728 84.539 Shaded wet ledges, sandstone cliffs, “rockhouses” (Fernald) King & H. Rob. Anemone lancifolia Pursh Ranunculaceae 164761 36.171 80.675 Damp rich woods Apios priceana B.L. Rob. Fabaceae 48167 35.295 87.156 Rocky limestone woods Astilbe biternata (Vent.) Britton Saxifragaceae 90349 35.968 83.680 Rich woods, north-facing banks & seeps Boechera perstellata (E.L. Braun) Brassicaceae 6767 36.855 85.985 Calcareous bluffs, wooded hillsides Al-Shehbaz Botrychium mormo W.H. Wagner Ophioglossaceae 130069 46.635 91.286 Rich basswood & sugar maple forest Boykinia aconitifolia Nuttall Saxifragaceae 71430 36.256 83.296 Moist woodland, water edges Cardamine ﬂ agellifera Brassicaceae 29109 35.852 82.736 Moist wooded slopes, ravines, seeps O.E. Schulz Cardamine micranthera Rollins Brassicaceae 3770 36.412 80.239 Moist woods, along streams & seeps Carex acidicola Naczi Cyperaceae 11721 32.576 85.805 Dry to mesic deciduous forest Carex austrocaroliniana L.H. Cyperaceae 58524 35.339 84.677 Rich moist deciduous and mixed forest Bailey 110/23/2013 5:19:51 PM

0 (Continued) / 2 3 / 2 0 1 3

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5 (Continued) 6 _ c 2 1 . i Species Family Range Area (km 2 ) Range Centroid Range Centroid Habitat Description n d d

Latitude (º) Longitude (º)

4 8 2 Carex basiantha Steudel Cyperaceae 153979 31.691 89.115 Mesic to wet-mesic deciduous forests Carex biltmoreana Mackenzie Cyperaceae 20185 35.001 82.627 Rocky woods, moist ledges, granite balds Carex brysonii Naczi Cyperaceae 8677 33.541 87.709 Mesic deciduous forest, slopes above streams Carex impressinervia Bryson Kral Cyperaceae 21699 32.604 86.166 Mesic deciduous forest, slopes above streams & Manhart Carex latebracteata Waterfall Cyperaceae 22384 34.489 94.140 Steep shaded slopes, mesic to dry-mesic forest Carex manhartii Bryson Cyperaceae 26810 35.654 82.765 Moist deciduous and mixed forest Carex ouachitana Kral Manhart Cyperaceae 19719 34.709 93.815 Mesic, dry-mesic rocky deciduous or mixed forest & Bryson Carex picta Steudel Cyperaceae 106445 35.079 86.973 Forests & forest openings Carex pigra Naczi Cyperaceae 33105 34.541 85.530 Mesic to wet-mesic deciduous forests Carex purpurifera Mack. Cyperaceae 91720 36.981 84.201 Moist deciduous forests, often near limestone ledges Carex radfordii Gaddy Cyperaceae 6738 34.908 82.832 Moist deciduous forests on calcareous soil Carex roanensis F.J. Herm. Cyperaceae 25391 37.632 80.928 Rich moist soil under beech trees Carex socialis Mohlenbr. & Cyperaceae 138778 34.543 87.543 Lowland deciduous forests, clay soils Schwegm. Carex superata Naczi, Reznicek Cyperaceae 56149 33.926 84.342 Moist to dry-mesic open deciduous forests, & B.A. Ford ravines Carex thornei Naczi Cyperaceae 12881 31.586 84.898 Mesic deciduous forests, slopes & ﬂ oodplains 110/23/2013 5:19:51 PM

0 Carex timida Naczi & B.A. Ford Cyperaceae 30531 37.221 87.338 Mesic deciduous or mixed woods, calcareous soil / 2 3 /

2 Chelone lyonii Pursh Scrophulariaceae 48756 35.430 83.152 Rich coves, stream banks 0 1 3

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6 Collinsonia tuberosa Michx. Lamiaceae 156125 33.890 84.845 Moist woods, calcareous soils _ c 2

1 Collinsonia verticillata B a l d w . L a m i a c e a e 1 0 4 7 9 3 3 4 . 6 4 0 8 3 . 5 5 8 W o o d e d s l o p e s , l o w w o o d s . i n d d Corallorhiza bentleyi Orchidaceae 4972 37.931 80.285 Deciduous forest & disturbed forest edges

4 8

3 Freudenstein Coreopsis delphiniifolia Lam. Asteraceae 57448 33.195 81.577 Woodlands, thickets & swamps Coreopsis latifolia Michx. Asteraceae 37579 35.293 82.818 Shaded slopes in rich moist woods Coreopsis pulchra Boynt. Asteraceae 15001 33.938 86.160 Forest openings, outcrops Croomia pauciﬂ ora (Nutt.) Torr. Stemonaceae 118397 32.445 86.422 Mesic wooded slopes & bottoms, circumneutral soils Cymophyllus fraserianus (Ker Cyperaceae 161869 37.209 81.412 Rich mesic shaded slopes in deciduous or mixed

Gawl.) Kartesz & Gandhi forest OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN Delphinium exaltatum Aiton Ranunculaceae 150440 38.378 80.962 Rocky slopes in rich woods or barrens, calcareous soil Delphinium newtonianum D.M. Ranunculaceae 15456 35.210 93.320 Slopes in deciduous forest Moore Desmodium humifusum (Muhl. Fabaceae 60252 41.435 73.884 Dry woods, sandy soils Ex Bigelow) Beck Desmodium ochroleucum M.A. Fabaceae 68327 36.163 82.393 Dry open woods, sandy or rocky soils Curtis ex Canby Dicentra eximia (Ker Gawl.) Torr. Fumariaceae 113533 38.533 80.477 Dry to moist rocky mountain woods, cliffs & crevices Diphylleia cymosa Michx. Berberidaceae 30681 35.653 82.732 Moist slopes, seeps & stream banks in deciduous forest (Continued) 110/23/2013 5:19:52 PM 0 / 2 3 / 2 0 1 3

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5 (Continued) 6 _ c 2 1 . i Species Family Range Area (km 2 ) Range Centroid Range Centroid Habitat Description n d d

Latitude (º) Longitude (º)

4 8 4 Dodecatheon amethystinum Primulaceae 77954 41.491 88.342 Moist hillsides & limestone cliffs in deciduous (Fassett) Fassett forest Dodecatheon frenchii (Vasey) Primulaceae 33569 36.935 88.144 Moist shaded ﬂ ats in woods under cliffs, near Rydb. streams Draba ramosissima Desv. Brassicaceae 189985 36.897 82.298 Rocky wooded areas, limestone cliffs, shale barrens Elymus svensonii Church Poaceae 15901 36.556 85.829 Woods on limestone bluffs, slopes & ledges Erythronium propullans A. Gray Liliaceae 8469 44.329 92.825 Mesic ﬂ oodplain woods Eupatorium godfreyanum Asteraceae 206635 37.821 80.203 Woods and disturbed open sites, forest edges Cronquist Euphorbia mercurialina Michx. Euphorbiaceae 154614 34.917 84.759 Rich soil on wooded slopes, ravines Euphorbia purpurea (Raf.) Euphorbiaceae 65719 38.408 79.312 Dry or moist woods Fernald Eurybia furcata (Burgess) Asteraceae 144841 41.253 89.021 North-facing slopes, moist deciduous woods G.L. Nesom Eurybia mirabilis (Torr. & Asteraceae 36392 34.616 81.526 Deciduous & mixed woods, slopes or alluvial A. Gray) G.L. Nesom plains Eutrochium steelei (E.E. Lamont) Asteraceae 53807 36.108 82.830 Open woods, gravelly banks, thickets E.E. Lamont Gentiana decora Pollard Gentianaceae 147285 36.058 82.497 Wooded slopes, coves, streambanks

110/23/2013 5:19:52 PM Geum geniculatum Michx. Rosaceae 4251 36.105 81.832 Balds and wooded coves at high elevation 0 / 2 3

/ Gymnocarpium appalachianum Dryopteridaceae 63167 39.360 79.509 Maple-birch-hemlock woods, tallus w/ cold air 2 0 1 3

Pryor & Hauﬂ er seepage

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6 Helianthus glaucophyllus Asteraceae 31974 34.361 84.283 Moist forests, woodland edges _ c 2

1 D.M. Sm. . i n d d Heuchera longiﬂ ora Rydb. Saxifragaceae 120569 36.636 83.826 Rich woods and roadcuts over limestone

4 8

5 Heuchera pubescens Pursh Saxifragaceae 223208 37.923 81.327 Shaded circumneutral rock outcroppings in woods Hexastylis contracta Blomquist Aristolochiaceae 35994 36.444 83.544 Acid soils in deciduous woods Hexastylis heterophylla (Ashe) Aristolochiaceae 220495 35.921 82.539 Deciduous & mixed forests Small Hexastylis lewisii (Fernald) Aristolochiaceae 84708 36.178 78.893 Upland & lowland forests, ﬂ oodplains Blomquist & Oosting

Hexastylis minor (Ashe) Aristolochiaceae 118466 36.385 79.880 Slopes & bluffs along streams in deciduous woods OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN Blomquist Hexastylis naniﬂ ora Aristolochiaceae 13875 35.342 81.773 Acidic soils on bluffs & ravines in deciduous Blomquist woods Hexastylis rhombiformis Gaddy Aristolochiaceae 5571 35.381 82.665 Deciduous woods on sandy river bluffs, ravines Houstonia serpyllifolia Michx. Rubiaceae 68074 35.968 82.651 Rich woods, stream margins, road cuts, pastures Hydrophyllum brownei Kral & Hydrophyllaceae 14729 34.463 93.601 Rich deciduous forests V.M. Bates Liatris gholsonii L.C. Anderson Asteraceae 3622 30.303 84.994 Slopes in deciduous woods, open xeric woods Lilium grayi S. Watson Liliaceae 27634 37.260 80.417 Moist forests, openings, bogs, seeps & wet meadows Listera smallii Wiegand Orchidaceae 137443 37.341 81.060 Damp humus in shady forests, under Rhododendron (Continued) 110/23/2013 5:19:52 PM 0 / 2 3 / 2 0 1 3

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5 (Continued) 6 _ c 2 1 . i Species Family Range Area (km 2 ) Range Centroid Range Centroid Habitat Description n d d

Latitude (º) Longitude (º)

4 8 6 Lysimachia tonsa (Alph. Wood) Primulaceae 188779 35.454 83.082 Moist hardwood forests, pine-oak woods, bluffs Alph. Wood ex Pax & R. Knuth Matelea alabamensis (Vail) Asclepiadaceae 12680 30.963 84.563 Slopes in deciduous forest Woodson Matelea baldwyniana (Sweet) Asclepiadaceae 91821 34.289 90.614 Open rocky woods, thickets Woodson Matelea ﬂ avidula (Chapm.) Asclepiadaceae 28298 32.215 83.229 Forested slopes & alluvial woods Woodson Meehania cordata (Nutt.) Britton Lamiaceae 280290 38.502 81.341 Rich mountain woods Monotropsis odorata Schwein. Monotropaceae 117575 35.895 81.484 Mixed deciduous or coniferous forests Ex Elliott Napaea dioica L. Malvaceae 144997 41.233 87.559 Moist alluvial woods Onosmodium decipiens J. Allison Boraginaceae 1595 32.997 87.124 Dolomite outcrops in rocky woods & glades Orbexilum onobrychis (Nutt.) Fabaceae 211639 36.834 85.907 Open woods, prairies Rydb. Oxalis illinoiensis Schwegm. Oxalidaceae 18788 37.076 86.559 Mesic to dry-mesic forests Penstemon deamii Pennell Scrophulariaceae 14461 38.374 87.972 Moist open woods, prairies Penstemon smalli A. Heller Scrophulariaceae 67622 35.223 84.115 Woodlands, cliffs, banks & forest edges Penstemon tenuis Small Scrophulariaceae 226272 32.491 92.333 Wet woodland soils, bottomlands Phacelia covillei S. Watson Hydrophyllaceae 18121 37.690 78.988 Rich soil of ﬂ oodplains & alluvial woods 110/23/2013 5:19:52 PM

0 Phacelia ﬁ mbriata Michx. Hydrophyllaceae 18475 35.225 84.269 Streambanks and alluvial woods / 2 3 /

2 Phacelia gilioides Brand Hydrophyllaceae 203809 36.851 93.099 Woodland openings, low rich woods, forest edges 0 1 3

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P M 487.1 ooxfordhb-9780199837656_c21.indd 487 x f o r d h b - 9 7 8 0 1 9 9 8 3 7 6 5

6 Phacelia ranunculacea (Nutt.) Hydrophyllaceae 86316 36.495 89.531 Mesic alluvial forests _ c 2

1 Constance . i n d d Platanthera integrilabia (Torr.) Orchidaceae 71905 34.626 85.345 Wet wooded ﬂ ats, seeps, wetlands

4 8

7 Polymnia cossatotensis Pittman Asteraceae 4264 34.518 93.949 Upland rocky woods & tallus, chert outcrops & V.M. Bates Polymnia laevigata Beadle Asteraceae 32810 34.956 86.666 Damp shaded sites, calcareous soils Prenanthes crepidinea Michx. Asteraceae 214423 39.023 87.680 Moist rich deciduous woods, thickets, prairies Prenanthes roanensis Asteraceae 64373 36.042 82.453 Spruce-hardwood forests, wooded slopes & balds (Chickering) Chickering Prosartes maculata (Buckley) Liliaceae 133520 36.723 83.839 Rich moist deciduous woods, slopes & ravines

A. Gray OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN Pycnanthemum beadlei (Small) Lamiaceae 9218 36.036 82.318 Forests, woodland borders Fernald Pycnanthemum curvipes Lamiaceae 11002 35.505 83.566 Dry rocky woodlands, rock outcrops (Greene) E. Grant & Epling Pycnanthemum loomisii Nutt. Lamiaceae 167430 36.288 83.308 Forests, woodland borders Pycnanthemum montanum Lamiaceae 60612 36.036 82.721 Balds, woodlands, forests & forest edges Michx. Pycnanthemum pycnanthemoides Lamiaceae 224378 35.977 83.790 Forests, woodland borders (Leavenworth) Fernald Pycnanthemum torrei Benth. Lamiaceae 118144 38.117 80.876 Dry rocky woodlands Ranunculus allegheniensis Britton Ranunculaceae 228781 39.716 78.869 Moist or dry woods, pastures Ranunculus harveyi (A. Gray) Ranunculaceae 181241 35.757 90.860 Acid soils on rocky wooded slopes, ridges, open Britton areas 110/23/2013 5:19:52 PM 0 / 2

3 (Continued) / 2 0 1 3

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P M 488.1 ooxfordhb-9780199837656_c21.indd 488 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9 9 8 3 7 6

5 (Continued) 6 _ c 2 1 . i Species Family Range Area (km 2 ) Range Centroid Range Centroid Habitat Description n d d

Latitude (º) Longitude (º)

4 8 8 Rudbeckia heliopsidis Torr. & Asteraceae 30072 34.134 82.963 Mesic to wet woodlands, meadows A. Gray Ruellia purshiana Fernald Acanthaceae 96854 35.755 82.650 Dry woodlands over calcareous rock Rugelia nudicaulis Shuttlw. ex Asteraceae 8435 35.416 83.423 High elevation spruce-fi r & northern hardwood Chapm. forest Salvia urticifolia L. Lamiaceae 222452 35.074 83.547 Rocky woodlands on circumneutral soils Scirpus fl accidifolius Cyperaceae 5145 36.680 77.301 Wooded bottomlands (Fernald) Schuyler Scutellaria arguta Buckley Lamiaceae 8601 37.422 82.573 Mesic woods and boulderfi elds at high elevation Scutellaria montana Chapm. Lamiaceae 11465 35.125 84.912 Open deciduous woods on mesic soil Scutellaria pseudoserrata Epling Lamiaceae 25289 34.273 85.258 Rich rocky forests Scutellaria saxatilis Riddell Lamiaceae 134303 37.039 82.940 Rocky forests, moist cliffs Scutellaria serrata Andrews Lamiaceae 160329 37.415 81.183 Rich deciduous forests Sedum glaucophyllum R.T. Crassulaceae 70801 36.266 80.519 Shaded cliffs, rocky slopes Clausen Shortia galacifolia Torr. & Diapensiaceae 16960 35.255 82.715 Moist forest slopes & stream banks in deep shade A. Gray Silene catesbaei Walter Caryophyllaceae 13101 32.253 84.080 Mesic deciduous forests along streams or slopes Silene nivea (Nutt.) Muhl. Ex Caryophyllaceae 278558 40.581 86.336 Rocky or fl ood-scoured alluvial woodlands Otth 110/23/2013 5:19:52 PM

0 Silene ovata Pursh Caryophyllaceae 94821 33.975 86.367 Woodlands & forests on circumneutral soil / 2 3 /

2 Silphium brachiatum Gattinger Asteraceae 14651 35.098 86.501 Open forests on calcareous soil, roadcuts 0 1 3

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P M 489.1 ooxfordhb-9780199837656_c21.indd 489 x f o r d h b - 9 7 8 0 1 9 9 8 3 7 6 5

6 Silphium wasiotense M. Medley Asteraceae 7680 36.852 83.588 Dry open sites in mesic forests _ c 2

1 Sisyrinchium dichotomum E.P. Iridaceae 6560 35.282 82.132 Dry to mesic oak-hickory forests . i n d d Bicknell

4 8

9 Solidago arenicola B,R, Keener Asteraceae 4422 35.138 85.582 Mesic woods in deep sandy alluvium & Kral Solidago auriculata Shuttlw. ex Asteraceae 132792 32.712 88.619 Rocky wooded slopes, alluvial soils S.F. Blake Solidago brachyphylla Chapman Asteraceae 50576 32.068 84.767 Open woodlands, bluff forests Solidago buckleyi Torrey & Asteraceae 46291 38.199 89.896 Open oak woods on ridges, slopes & bluffs A. Gray

Solidago curtisii Torrey & A. Gray Asteraceae 199625 36.086 83.685 Shaded mesic woods & thickets OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN Solidago drummondii Torrey & Asteraceae 5268 35.384 93.380 Limestone ledges & bluffs in rocky woods A. Gray Solidago faucibus Wieboldt Asteraceae 27460 36.378 83.214 Mesic deciduous forests & hardwood-hemlock Solidago ﬂ accidifolia Small Asteraceae 65118 34.581 85.173 Mesic woods & clearings Solidago lancifolia (Torrey & Asteraceae 4481 36.214 82.168 Rich woods, mountain slopes, road embankments A. Gray) Chapman Solidago ouachitensis C.E.S. Asteraceae 12521 34.697 94.053 Woods on north-facing slopes Taylor & R.J. Taylor Solidago roanensis Porter Asteraceae 166235 36.793 82.307 Forests, woodlands, roadbanks, edges of mtn balds Solidago sphacelata Raﬁ nesque Asteraceae 210540 36.501 84.152 Open woods & rocky places, calcareous soils Spigelia loganioides (Torr. & Loganiaceae 13586 29.101 82.055 Wet calcareous hammocks & woods A. Gray ex Endl. & Fenzl) A. DC. 110/23/2013 5:19:53 PM 0 / 2 3 /

2 (Continued) 0 1 3

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P M 490.1 ooxfordhb-9780199837656_c21.indd 490 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9 9 8 3 7 6

5 (Continued) 6 _ c 2 1 . i Species Family Range Area (km 2 ) Range Centroid Range Centroid Habitat Description n d d

Latitude (º) Longitude (º)

4 9 0 Stachys clingmanii Small Lamiaceae 22374 37.965 86.496 Cove forests & boulderﬁ elds at high elevation Stachys cordata Riddell Lamiaceae 205560 37.270 83.244 Moist forests, alluvial soils or over calcareous rock Stachys eplingii J.B. Nelson Lamiaceae 31266 36.409 82.080 Mtn woods, mesic forests, bogs & wet meadows Stachys iltisii J.B. Nelson Lamiaceae 29848 35.582 93.084 Rich soil in open upland woods Stachys latidens Small ex Britton Lamiaceae 55908 36.708 81.645 Mesic forests in coves, forest edges Stellaria corei Shinners Caryophyllaceae 127557 37.891 84.276 Mesic cove forests & seeps at mid- to high-elevation Symphyotrichum anomalum Asteraceae 245731 37.523 91.701 Rocky open deciduous woods, dry ridges, cliffs, (Engelm.) G.L. Neson bluffs Symphyotrichum phlogifolium Asteraceae 229281 37.520 82.037 Rich mesic mixed hardwood forests, roadsides (Muhl. ex Willd.) G.L. Nesom Symphyotrichum retroﬂ exum Asteraceae 64470 34.405 83.084 Moist woodlands, meadows, open pine or oak (Lindl. ex DC.) G.L. Nesom woods Synandra hispidula (Michx.) Lamiaceae 121940 38.105 84.694 Rich mesic woods Britton Thalictrum clavatum DC. Ranunculaceae 105376 36.293 83.545 Rich moist woods, cliffs, seeps, stream banks Thalictrum coriaceum (Britton) Ranunculaceae 92381 37.382 81.760 Rocky or mesic open deciduous woods, thickets Small Thalictrum debile Buckley Ranunculaceae 28466 33.444 87.312 Rich, rocky woods on limestone, wet alluvial soil Thalictrum macrostylum Small & Ranunculaceae 81502 34.951 80.734 Rich wooded slopes, cliffs, swamp forests,

110/23/2013 5:19:53 PM A. Heller meadows 0 / 2 3 /

2 Thalictrum mirabile Small Ranunculaceae 15112 35.523 86.089 Moist bluffs, wet sandstone cliffs, sinks 0 1 3

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P M 491.1 ooxfordhb-9780199837656_c21.indd 491 x f o r d h b - 9 7 8 0 1 9 9 8 3 7 6 5

6 Thaspium pinnatiﬁ dum Apiaceae 15560 36.586 84.752 Forests & woodlands over calcareous rock _ c 2

1 (Buckley) A. Gray . i n d d Thermopsis fraxinifolia Nutt. ex Fabaceae 42342 34.942 83.022 Dry slopes, ridges & clearings

4 9

1 M.A. Curtis Thermopsis mollis (Michx.) M.A. Fabaceae 80831 35.746 82.320 Dry slopes, open woods & clearings Curtis ex A. Gray Thermopsis villosa (Walter) Fabaceae 37056 35.877 82.885 Mesic forest openings, ﬂ oodplains & roadbanks Fernald & B.G. Schub. Tragia cordata Michx. Euphorbiaceae 172584 33.632 89.576 Rich woods over limestone, rocky hillsides Trifolium stoloniferum Muhl. Fabaceae 102702 38.140 87.699 Moist disturbed forests, streams, open woods,

ex Eaton lawns OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN Trillium decipiens J.D. Freeman Melanthiaceae 41712 31.828 84.671 Rich woods & river bluffs in mixed deciduous forests Trillium decumbens Harbison Melanthiaceae 38956 33.870 85.463 Rocky slopes in open deciduous woodlands Trillium discolor Wray ex Hook. Melanthiaceae 15733 34.339 82.551 Forested slopes & stream banks Trillium foetidissimum J.D. Melanthiaceae 71867 31.105 92.228 Rich woods on river bluffs, ﬂ oodplains, roadsides Freeman Trillium gracile J.D. Freeman Melanthiaceae 47927 31.009 94.179 Mature pine & hardwood forests, slopes near streams Trillium lancifolium Raf. Melanthiaceae 68672 33.317 85.249 Floodplain forests, rocky upland woods & thickets Trillium ludovicianum Harbison Melanthiaceae 61448 31.552 92.487 Mixed deciduous ﬂ oodplain woods & adj. slopes Trillium luteum (Muhl.) Melanthiaceae 71780 34.354 83.826 Rich deciduous forest & open woods, calcareous Harbison soils (Continued) 110/23/2013 5:19:53 PM 0 / 2 3 / 2 0 1 3

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P M 492.1 ooxfordhb-9780199837656_c21.indd 492 x f o OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN r d h b - 9 7 8 0 1 9 9 8 3 7 6

5 (Continued) 6 _ c 2 1 . i Species Family Range Area (km 2 ) Range Centroid Range Centroid Habitat Description n d d

Latitude (º) Longitude (º)

4 9 2 Trillium maculatum Raf. Melanthiaceae 92959 31.660 83.938 Rich mesic forests, river banks & bluffs, fl oodplains Trillium oostingii Gaddy Melanthiaceae 1742 34.352 80.583 Rich bottomland forests Trillium persistens Duncan Melanthiaceae 2970 34.755 83.198 Mixed deciduous & pine woodlands, stream fl ats Trillium pusillum Michx. Melanthiaceae 92822 36.115 85.434 Dry to mesic forests, along streams, swampy woods Trillium reliquum J.D. Freeman Melanthiaceae 26511 32.315 84.163 Rich mixed forest, slopes, bluffs & stream fl ats Trillium rugelii Rendle Melanthiaceae 91092 35.133 83.850 Rich deciduous forests, calcareous or mafi c bedrock Trillium simile Gleanon Melanthiaceae 26375 35.307 83.421 Forested coves, slopes & seeps with rich soil Trillium stamineum Harbison Melanthiaceae 97060 33.645 87.520 Upland deciduous forest over limestone, fl oodplains Trillium sulcatum Patrick Melanthiaceae 99413 38.779 80.968 Coves & moist slopes, rich mesic woodlands Trillium underwoodii Small Melanthiaceae 75918 31.587 85.220 Dry to mesic rich deciduous forests, stream edges Trillium vaseyi Harbison Melanthiaceae 36463 33.210 84.492 Steep wooded slopes, rich coves & ravines Trillium viride Beck Melanthiaceae 41016 38.471 90.572 Rich woods, bluffs & rocky hillsides Trillium viridescens Nutt. Melanthiaceae 120456 34.685 93.618 Rich deciduous forests, bluffs & fl oodplains Uvularia fl oridana Chapm. Liliaceae 48386 32.203 84.840 Rich hardwood forests, fl oodplains & moist ravines Valeriana paucifl ora Michx. Valerianaceae 213927 38.998 84.412 Rich mesic woods 110/23/2013 5:19:53 PM 0 / 2 3 / 2 0 1 3

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P M ooxfordhb-9780199837656_c21.indd 493 x f o r d h b - 9 7 8 0 1 9 9 8 3 7 6 5

6 Veratrum latifolium (Desr.) Liliaceae 164689 36.651 81.255 Moist to dry forests _ c 2

1 Zomlefer . i n d d Veratrum parviﬂ orum Michx. Liliaceae 110403 35.900 83.904 Moist wooded slopes, dry forests

4 9

3 Veratrum woodii J.W. Robbins ex Liliaceae 183745 37.339 87.774 Rich woods on circumneutral soil Alph. Wood Vernonia arkansana DC. Asteraceae 188301 39.133 93.390 Low woods, streambanks, roadsides Viola tripartita Elliott Violaceae 214221 35.650 83.407 Rich woods, moist slopes, bottomlands Viola villosa Walter Violaceae 251059 33.182 88.085 Moist sandy or rocky soil, hardwood hammocks Waldsteinia lobata (Baldw.) Torr. Rosaceae 16248 33.792 83.900 Forests, streambanks & A. Gray

Xerophyllum asphodeloides (L.) Liliaceae 86181 36.641 81.026 Forests on dry ridges & slopes, pine barrens OUP UNCORRECTEDPROOF –FIRSTPROOFS, Wed Oct232013,NEWGEN Nutt. Zizia trifoliata (Michx.) Fernald Apiaceae 228512 34.441 83.091 Mesic forest, woodlands, forest edges 110/23/2013 5:19:53 PM 0 / 2 3 / 2 0 1 3

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P M OUP UNCORRECTED PROOF – FIRSTPROOFS, Wed Oct 23 2013, NEWGEN

ooxfordhb-9780199837656_c21.inddxfordhb-9780199837656_c21.indd 449494 110/23/20130/23/2013 55:19:53:19:53 PPMM REFERENCES

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