1 North America Defies a Global Trend of Reduced Flowering Plant Diversity in Extreme Environments

2 Robert D. Edwards1, Chase M. Mason2, Travis W. Nauman3, Martin B. Goldhaber4, Elisabeth N. Bui5, Vicki

3 A. Funk1, James A. Thompson6, Pamela S. Soltis7,8, Jennifer M. Cartwright9

1
4 U.S. National Herbarium, Department of Botany, National Museum of Natural History, Smithsonian

5 Institution, Washington, D.C. 20013-7012, USA.

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6 University of Central Florida, Orlando, FL 32816, USA.

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7 U.S. Geological Survey, Southwest Biological Science Center, Moab, UT 84532, USA.

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8 U.S. Geological Survey, Denver, CO 80225, USA.

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9 Division of Land and Water, CSIRO, Canberra, ACT, Australia.

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Division of Plant and Soil Sciences, West Virginia University, Morgantown, WV 26506, USA.
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11 Florida Museum of Natural History, University of Florida, Gainesville, FL 32611, USA.
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12 Biodiversity Institute, University of Florida, Gainesville, FL 32611, USA.

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13 U.S. Geological Survey, Lower Mississippi-Gulf Water Science Center, Nashville, 37211 TN, USA.
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15 Abstract

16 It is generally assumed that more extreme environments are less habitable for a majority of organisms,

17 leading to a decrease in diversity. However, as definitions of extreme are typically based on particular

18 organisms’ tolerances and may not be applicable across biodiversity, this assumption has not been well

19 tested. Here we use the upper and lower ranges of environmental variables to derive a set of 30

20 extreme stress layers that fall within nine extreme stress categories, each expected to affect the survival
21 of terrestrial organisms. We use these to show that while flowering plant species richness globally

22 decreases as the environment becomes more extreme, the reverse is true in North America where

23 increased species richness is associated with extremeness in the arid, high energy environments of the

24 south and west of the continent. Data for the Compositae family of plants (asters) in North America

25 support this finding and suggest that the diversity of this group alone in challenging environments

26 contributes much to this response. Our results show the utility of defining the extremeness of an

27 environment agnostic of organismal tolerance. Hostile environments generally do not promote plant

28 diversity, but by identifying these environments we can detect lineages that prosper under such

29 conditions.

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31 Introduction ie
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32 Understanding the range and distribution of extreme conditions and hostile environments is particularly

33 important under a changing global climate. Already, increases in the frequency and intensity of weather
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34 and climate events are evident1,2, and there is an understanding that changing conditions may push

35 many organisms towards, or beyond, their physiological limits3. Identifying regions where extreme
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36 conditions already exist is important for conservation not only because these environments are at an

37 elevated risk of losing biodiversity as organismal physiological limits are surpassed, but also because

38 they support organisms that already have adaptive traits and genetic potential for tolerating changes

39 elsewhere4. Yet defining what makes an environment extreme is not straightforward and faces two

40 primary challenges: firstly, identifying a threshold above or below which any particular environmental

41 variable can be considered extreme is often subjective given that what to one organism is extreme may

42 be optimal and normal to another5; and secondly, that multiple variables may contribute to the severity

43 of an environment.
44 We can address the first of these challenges by observing that while organismal tolerances of any

45 environment are to some degree idiosyncratic, for terrestrial organisms the comfortable envelope for

46 survival and reproduction typically exists towards the middle of the range of most environmental

47 variables (as experienced at ground level)6. Stated inversely, organismal tolerances tend to fall off at the

48 upper and/or lower limits of a variable’s range (e.g., extreme cold or heat)7. As such, an environmental

49 extreme can be defined as a statistically rare occurrence well outside the bounds of normal variability6,8.

50 This means that we can objectively define “extreme” as a small fraction of a variable’s range in the

51 distribution tail near its upper or lower limit, and subsequently we can delimit the land areas where

52 these values are found. By defining extremes in this way - abiotically based on environmental values

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53 rather than the perceived preferences of organisms - we can directly test the response of biotic

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processes, such as diversity, organismal traits, body size, etc.

The second challenge is to consider how multiple variables contribute to the extremeness of an
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56 environment. Above we have defined the extremity of single variables; however, many variables may

57 have similar physiological effects on organisms. For example, high topographic wetness, high annual
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58 precipitation, high precipitation of the wettest month, and high mean annual humidity can all represent

59 high moisture extremes. By combining these layers into a single class (implicitly assuming a flat prior of
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60 equal weight), we allow questions to be asked of the overarching mechanism of high moisture stress

61 (Fig. 1).

62 From an evolutionary perspective, it is expected that adaptation of a lineage to an environment with a

63 single or few such physiological stressors is less difficult than adaptation to many different physiological

64 stressors simultaneously. Trade-offs and the multifaceted effects of many stressors make it increasingly

65 difficult for a lineage to develop a viable ecological strategy as the number of co-occurring pressures

66 increases. For example, while some traits can confer adaptive advantage across a number of stressful

67 factors (the C4 photosynthetic pathway reduces wasteful photorespiration under high light, heat,
68 drought, and salinity9,10; osmotic adjustment provides resistance to low water potentials under salinity,

69 drought, and frost11), such strategies generally involve concessions against the optimization of other

70 functions12–14, and may come at the expense of resistance to other stressors15,16. Indeed, such is the

71 genetic architecture of many quantitative traits that selection conferring resistance to one particular

72 stressor may actually reduce resistance to others17,18. Given these factors, it is expected that it should be

73 difficult for a lineage to deal with a larger number of simultaneous environmental stressors, making the

74 presence of multiple environmental extremes a substantial barrier to colonization and diversification.

75 In this study we define the extreme limits of 30 abiotic variables on a global scale, identify nine classes

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76 of physiological stress to which these belong, and test the hypothesis that as the number of overlapping

77 classes increases, there will be a decrease in the diversity of flowering plants globally, as well as for each

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continent in isolation. We also test the response of the largest North American family of flowering plants

(Compositae – asters) to environmental extremes across this continent.

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81 Results

82 Extreme layers and classes

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83 From climate and substrate data we have identified 30 variables (layers) for which the upper or lower

84 5% of values are expected to present extreme challenges to terrestrial plants. These 30 layers fall under

85 nine extreme classes: energy limitation, water limitation, nutrient limitation, heat stress, cold stress,

86 extreme variability, UV exposure, high moisture stress, and soil physical stress (Table S1). Two-thirds

87 (66%) of cells are subject to at least one of these extremes (Fig. 2). When translated into land area, no

88 1km2 cell contained more than 17 (of a possible 30) extreme layers, while only 412 cells (2.5%)

89 represented the maximum 9 extreme classes (Table S2). Grid cells with the highest values (14-17

90 extreme layers, or 9 extreme classes) were all restricted to the highest peaks of the Bolivian Andes.
91 There is strong geographic regionalization of extreme layer and class number with high numbers co-

92 occurring on the western margins of North America and South America, southern Africa, the Tibetan

93 Plateau, southeastern Australia and the Pilbara, all regions with relatively low flowering plant diversity

94 (Fig. S1). Low diversity is, however, also found in the presence of only a modest number of extreme

95 classes at high latitudes, as well as in Patagonia, Saharan Africa, the Middle East, the Taklamakan Desert

96 and southern and central Australia.

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98 Correlation of Extremeness with Flowering Plant Diversity

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99 We use global flowering plant diversity data to test the hypothesis that as the number of overlapping

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extreme layers or classes increases, plant diversity will decline. As expected at a global scale, plant

diversity generally is highest (significantly greater than random) when the number of extreme classes is
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102 low (zero or one) and decreases rapidly as the number of classes increases (Fig. 3A). This pattern is also

103 true for each of the individual stress classes when examined in isolation, except energy limitation where
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104 richness is significantly greater than expected when three stressors are present, and for UV exposure

105 where there is a strong and significant increase in richness as the number of stressors increases (Fig. 4).
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106 Note that due to the coarse scale of the available plant richness data (7800km2 polygons), extreme class

107 values derived from the centroids of these polygons failed to capture the rare cells with maximum

108 values (14-17 classes). Tests for possible effects of latitude in driving correlations between the number

109 of environmental extremes and species richness suggest little influence, with regression of the number

110 of extreme classes against latitude returning only weak correlations for all continents (R2 < 0.145; Fig.

111 S2). Similarly, analyses using residuals of diversity against latitude were qualitatively the same as those

112 using raw richness, although dampened (Fig. S3). Only those results using raw richness will be discussed

113 here.
114 At continental scales, Africa, Eurasia, and South America (Figs. 3B, C, E) generally exhibit similar patterns

115 to the global data; however, Australia and North America show different responses (Figs. 2D and F).

116 North American plants are less diverse than expected where few extreme classes co-occur (zero or one)

117 with increasing diversity up to five concurrent extreme classes. A decrease in diversity is only evident

118 when six or more extreme classes are overlapping. This holds true even with the inclusion of glaciated

119 areas of Greenland, although the signal is slightly dampened (Fig S4). However, the trend of greater

120 richness with increased number of extreme classes in North America is almost eliminated when

121 Compositae richness is subtracted from total plant richness (Fig 3H). When analyzed in isolation, North

122 American Compositae species numbers increase sharply in response to a higher number of overlapping

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123 extreme classes and are significantly higher than expected by chance except where zero extreme classes

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are present (Fig. 3G). Geographically this translates to greater species richness in the more extreme USA

southwest and western Mexico (Fig. 5).

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126 For Australia, flowering plant diversity is neither greater nor less than expected by chance except where

127 there are no extreme classes or the highest number of co-occurring classes (eight) are found; here
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128 diversity is greater than expected.

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129 Results for the frequency of different class combinations across continents (Fig. S5) show considerable

130 variation, with some unique combinations, especially for higher numbers of classes (seven or eight).

131 There are, however, very few combinations unique to North America.

132 Analyses within individual extreme classes show that only two, energy limitation and high moisture, are

133 correlated with increased plant diversity across all continents (Fig. S6). Nutrient limitation, extreme

134 variability, soil physical stress, and water limitation are correlated with decreased richness on all

135 continents. Australia and North America are the only continents to show increased richness with greater
136 UV exposure, with Australia also showing increased richness for elevated water limitation, and North

137 America for heat stress.

138

139 Discussion

140 Summarizing extreme values across biologically relevant environmental variables is an important

141 conceptual step in being able to explore the responses of organisms to extreme conditions. We find that

142 while two thirds of the Earth’s land area, most of which is not considered environmentally challenging, is

143 subject to at least one extreme class, additional extremes occur in regions typically considered harsh. In

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144 some instances such as eastern Saharan Africa, the Middle East, the Taklamakan Desert, central

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Australia, and arctic tundra, there are just one or two overlapping classes. These are typically water

limitation/heat stress/UV exposure in hot deserts or water limitation/cold stress at high latitudes. Other
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147 areas with a greater number of co-occurring extreme classes are also known to be unfavorable for plant

148 growth, such as high elevations in the Andes (the only place on Earth where the number of classes
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149 reached the maximum of nine), the southwest of the USA and western Mexico, the Atacama desert,

150 western Sahara, the Middle East, Tibetan Plateau, and glaciated areas of Greenland. However, whether
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151 the cumulative effect of additional stressors results in measurably more inhospitable environments is

152 not immediately obvious. To address this, we use extreme class layers and plant occurrence data to

153 show that global flowering plant diversity decreases in areas with a higher number of overlapping

154 extreme classes, but that this trend is inverse in North America.

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156 Increased extremeness is associated with decreased species richness globally

157 At a global scale our data are consistent with the expectation that an increased number of overlapping

158 extreme stressors results in decreased flowering plant diversity. Notably there is a spike in diversity

159 above the null distribution for areas with a single extreme class, both globally and independently for the

160 continents of South America, Africa, and Eurasia. This indicates that while environments with multiple

161 extreme factors are challenging, the presence of only a single class of stressor may actually promote

162 species accumulation.

163 At continental scales, South America, Africa, and Eurasia display similar patterns to those found globally

164 with decreased diversity as the number of extreme classes increases (highly pronounced in South

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165 America); however, Australia and North America show different responses. Little significant variation in

166 diversity in Australia may reflect a long-term broadly stable and isolated landscape with a generally

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uniform fulfillment of niches19. An uptick in diversity where the most extreme classes overlap

corresponds to several well-known biodiversity hotspots (eastern and southwestern Australia20–22). This
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169 increase in diversity where conditions are most extreme is the same across continents (although only

170 significant in Australia) and suggests that the most stressful environments may actually promote species
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171 richness. The mechanism for this is unclear and should be further investigated.
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172 Extremeness in North America is associated with increased species richness

173 North America is the only continent to show a rise in species richness as the number of overlapping

174 extreme classes increases (for all but the two highest values of overlapping classes). This is consistent

175 with recent (re)colonization of mid-to-high latitudes subsequent to glaciation, resulting in low diversity

176 despite moderate contemporary environments in these regions23, contrasting with greater richness in

177 southern Mexico and the North American Coastal Plain where longer-term stability of environments has

178 been linked to increased endemism and diversity20,24–26. Only when the number of concurrent extreme

179 classes is highest (six and seven) in the deserts of southwestern USA and the Sierra Madres Occidental of
180 Mexico, does North American species richness decline (Fig. 3F and S5A). A similar signal is likely not seen

181 in the Eurasian data as a result of the Eurasian ice-sheet covering a much smaller proportion of the total

182 landmass in that region27.

183 In North American Compositae (a family of plants recognized as thriving in marginal environments),

184 species richness increases as the number of extreme classes increases. Indeed, our results for this group

185 of species in isolation (Fig. 3G) demonstrates a strong increase in species richness where more extreme

186 classes overlap. So strong is this relationship that when Compositae richness is subtracted from total

187 flowering plant richness for North America, the trend of greater richness with increased extremeness

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188 across all flowering plants is almost removed (Fig. 3H). This suggests that the Compositae are

189 responsible for driving much of the increased diversity in the challenging habitats of North America, and

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that an ability to thrive and diversify in extreme environments may at least partly explain why this is the

largest family of the flowering plants not only in North America, but globally.28
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192 That plant richness responds differently to extremes in North America could also be governed by which
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193 combinations of classes exist on that continent. Examination of the frequency of class combinations for

194 each continent (Fig. S5) does not suggest that this is the case, as while there is considerable variation in
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195 the combinations of classes across continents, few are unique to North America.

196 It is also possible that species richness in North America responds idiosyncratically to one or several

197 individual classes, driving the observed response. Examining how individual extremes affect species

198 richness on each continent (Fig. S6), we find generally similar patterns: plant diversity decreases as

199 extremeness increases, except for high-moisture extremes where diversity increases, and energy

200 limitation which has differing relations to diversity among the continents. The association of angiosperm

201 richness with water and energy availability are known phenomena29–31; however in North America we

202 also find that richness increases with greater extremeness in heat stress and UV exposure.
203 Conclusions

204 The data we present here summarize abiotically extreme areas in single geospatial layers for exploring

205 organismal response to extremeness across individual stressors as well as classes of environmental

206 stressors. We used these data to test two hypotheses and find that while global flowering plant richness

207 decreases as the environment becomes more extreme, the inverse is true in North America where

208 increased richness is associated with arid, high-energy environments of the south and west of the

209 continent. Much of this signal in North America is driven by the Compositae family, with our findings

210 suggesting that the diversity of this group alone in extreme environments influences patterns in the

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211 North American flora more generally.

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213 Methods
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214 Defining Extreme Thresholds

215 Extreme values of climatic and edaphic variables may represent environmental thresholds that influence
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216 speciation patterns and species richness7. A candidate set of 40 environmental variables was compiled

217 from WorldClim (19 climate variables32) and SoilGrids1km (21 substrate and topographic variables33).
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218 This was reduced to 24 by removing those that are highly autocorrelated. Each of the remaining

219 variables was identified as being either single or two-tailed (having a biological effect either at one or

220 both extreme ends), with two-tailed variables being separated into distinct high and low variables for a

221 final set of 30 derived variables. A standard extreme threshold of 95% was set for each variable, with the

222 values for ≥95th percentile calculated for high-end extremes, and ≤5th percentile calculated for low-end

223 extremes. While 95% is effectively arbitrary, it is a familiar significance cutoff when considering

224 distributions and likely conservative. The land area falling under these values was plotted with grid cells

225 at a resolution of 1km2 and these cells are provided as geotiffs for use in further research (Fig. S7). To
226 account for global heterogeneity in variable ranges, calculations were done for each continent (North

227 America, South America, Eurasia, Africa, and Australia) independently, and then these results were

228 combined. As continents largely represent biogeographic islands (and map closely to recognized

229 biogeographic realms34), delimiting relative extremes within these areas independently also

230 acknowledges that a majority of organisms within a continent are likely to have evolved in response to

231 the ranges of environmental variables in that region, rather than those at a global scale. To determine

232 where the highest overlap of extreme variable layers occurs, we generated a heatmap representing the

233 layer count for each cell (Fig. 1A). Variables were then grouped into nine classes according to the type of

234 stress they confer: energy limitation, water limitation, nutrient limitation, heat stress, cold stress,

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235 extreme variability, UV exposure, high moisture stress, and soil physical stress (Table S1). Some variables

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were attributable to multiple classes. As with the extreme layers, geotiffs for each individual class were

generated as well as a total extremeness layer where grid cell values were summed across extreme
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238 classes (Fig. 1B). Geotiffs of all individual and total extremeness layers are available via the Dryad Digital

239 Repository.
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241 Correlation of Extremes with Global Plant Diversity

242 To test the hypothesis that plant diversity is expected to decrease as the number of overlapping

243 extreme classes increases, global native vascular plant richness values (for the non-permanently iced

244 terrestrial Earth surface) were obtained from Ellis, Antill, & Kreft (2012) for 16,461 equal area polygons

245 (each 7800km2) using ArcMap v10.3 (ESRI, Redlands, CA). As much of Greenland is permanently iced and

246 potentially represents a not insignificant portion of cold and radiation extremes of North America, 238

247 polygons representing the glaciated area (unglaciated areas were already present) were added to the

248 above dataset and given a plant richness value of zero.

249 Within extreme classes, the constituent layers were flattened, resulting in a binary presence/absence

250 layer for each (the centroids for 25 polygons from the plant richness dataset fell outside the margins of

251 the extreme layers resulting in no data value for these points, and were excluded). The number of

252 extreme classes (0-9) present at the center of each of the above polygons was then also calculated.

253 Polygons were binned by the number of extreme classes present and the average vascular plant richness

254 calculated for each bin. From this we plotted average species richness against the number of extreme

255 classes. To determine whether the observed richness for each class was greater, less than, or no

256 different than that expected by chance, we generated a randomized distribution using the dplyr package

257 in R (R v3.5.1, R Core Team 2018). This was done by holding the value for the number of extreme classes

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258 constant for each cell and randomly shuffling the observed richness values 999 times. The average

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richness value for each number of extreme layers was calculated for each of the 999 permutations. The

observed average for each number of extreme layers was compared to this distribution. An observed
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261 value falling above or below the top or bottom 5% of the randomized values was considered significantly

262 higher or lower than expected by chance. To understand how the relationship between the number of
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263 co-occurring extreme classes and richness may vary at continental scales, these methods were repeated

264 for each continent in isolation. In order to determine how individual extreme classes influence plant
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265 diversity, analyses were also conducted for the layers within each class in isolation. As a number of

266 abiotic variables have been shown to be correlated with the Latitudinal Diversity Gradient (LDG) 30 the

267 above analyses were also run with the residuals from richness regressed against latitude to correct for

268 any association with latitude. Regressions for extremeness (number of extreme layers) against latitude

269 were also conducted for each continent to further explore a possible gradient effect. To explore the

270 number and distribution of different combinations of extreme classes, frequency plots were compared

271 for all combinations of 3-8 classes on each continent using the UpSetR package in R 36.

272 North American Compositae

273 Occurrence data for 3,116 North American Compositae species (representing 14 tribes; Greenland was

274 not included) were also aggregated from GBIF, iDigBio, and BISON, cleaned (Edwards et al., in prep), and

275 the species richness scores for 100 km by 100 km grid cells calculated using Biodiverse v2.137. The

276 number of extreme layers and classes at the center of each of these grid cells was also recorded. Using

277 these data, the same analyses were run as for the total flowering plant diversity above. To understand

278 what influence Compositae have in driving the response of total flowering plant diversity in North

279 America, we subtracted the richness values for the Compositae from the total flowering plant data for

280 each cell and re-ran the analyses.

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281

282 Data Availability

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Geotiffs for all layers of individual extreme variables and extreme classes as well as the spatial data used
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284 for the calculation of Compositae species richness will be available for general use in the Dryad Digital

285 Repository on publication.

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287 Code Availability

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288 R code for calculating percentiles available at: https://github.com/BortEdwards/Delimiting-numeric-and-

289 geographic-thresholds-for-environmental-variables)

290 R code for randomizing richness values by class available at:

291 https://github.com/BortEdwards/Randomizing-species-richness-by-class)

292

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371 Acknowledgements

372 This work was funded by a USGS John Wesley Powell Center for Analysis and Synthesis grant to E.B. and

373 M.G., with support from iDigBio (NSF grant DBI-1547229 to PSS) and the University of Florida, and in-

374 kind support from the Smithsonian National Museum of Natural History’s National Herbarium. This work
375 would not have been possible without the invaluable contributions of many herbaria and botanists

376 world-wide to digital databases. We also thank Dr Meghan Balk for helpful insights on this project, and

377 Dr Joe Miller and Dr Mike Crisp for comments on a draft manuscript. Any use of trade, firm, or produce

378 names is for descriptive purposes only and does not imply endorsement by the U.S. Government.

379

380 Author Contributions

381 E.B. and M.G. conceived the project. E.B., M.G., R.E., C.M., J.C., T.N., J.T., V.F., and P.S. all contributed to

382 variable selection and design of analyses. R.E., J.C., T.N. and J.T. compiled the data. R.E., C.M., and J.C.

w
383 performed analyses. R.E. wrote the manuscript.

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A
By Continent: Top 95th percentile of Mean Annual Humidity

95% 95%

95% 95% 95%

Individual variable layer: Top 95th percentile of Topographic Wetness Index

B
Individual variable layer: Top 95th percentile of Annual Precipitation

Individual variable layer: Top 95th percentile of Precipitation of Wettest Month

w
Individual variable layer: Top 95th percentile of Mean Annual Humidity

ie
ev
C
Class: High Moisture Extremes
R
In

Number of overlapping
extreme layers
4
3
2
1
0

Figure 1. Schematic representation of the method for the constructing extreme classes: (A) calculation of upper
or lower 5th percentiles for each continent individually and recombination into a single global layer;
(B) selection and stacking of layers belonging to each extreme class; (C) combining stack into a single extreme
class layer with number of overlapping extremes indicated as a heat map.
 A 14

12

10

ew
0

i
8
ev
6
R
4
In

0
Figure 2. Heat map showing the number of (A) overlapping extreme layers globally, and
(B) overlapping extreme classes globally. Color scale depicts the ranges of values visible at global
scales, however a very small number of grid cells in the Andes Mountains of South America
(16 or 17 extreme layers and 9 extreme classes) cannot be represented at this resolution. Areas
not evaluated are represented in dark gray. Schematic for the generation of extreme layers and
classes is outlined in Figure 1.
 A Global B Africa C Eurasia D Australia
2500 2000 1500

1500 2000

1500
1500 1000

Richness
Richness

Richness

Richness
1000
1000 1000

500
500

ew
0 500 0
0 1 2 3 4 5 6 7 8 0 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7 8
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

i
North American North American Flowering

ev
E South America F North America G Compositae H Plants Minus Compositae
1500 4500
5000
4000
4500

4000

3500 1000
R 500
3500

3000

2500
In
3000 Richness

Richness
Richness

Richness

2500 2000

2000 1500
500
1500 1000
1000
500
500
0 0
0 1 2 3 4 5 6 7 8 0 0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7 8 0 1 2 3 4 5 6 7 8
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes
Figure 3. Average richness for all flowering plants plotted against the number of extreme categories (A) globally, (B-F) for each continent independently, and (G)
for North American Compositae only. Randomizations of the data in blue, observed data line green if significantly greater or less than expected by chance (falling
outside ≥97.5%, ≤2.5% of randomizations), red if not significantly outside of random.
 Global Cold Stress Global Energy Limitation Global Extreme Variability
5000 5000 5000

4500 4500 4500

4000 4000 4000

3500 3500 3500

3000 3000 3000

Richness
Richness
Richness

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 0 0

0 1 2 3 0 1 2 3 0 1 2 3 4 5
Number of Classes Number of Classes Number of Classes

Global Heat Stress Global High Moisture Stress Global Nutrient Limitation
5000 5000 5000

4500 4500 4500

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4000 4000 4000

3500 3500 3500

3000 3000
ie 3000
Richness

Richness

Richness
2500 2500 2500
ev
2000 2000 2000

1500 1500 1500

1000 1000 1000

R

500 500 500

0 0 0
0 1 2 3 0 1 2 3 0 1 2 3 4 5
Number of Classes Number of Classes Number of Classes
In

Global Soil Physical Stress Global UV Exposure Global Water Limitation

5000 5000 5000

4500 4500 4500

4000 4000 4000

3500 3500 3500

3000 3000 3000

Richness

Richness

Richness

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 0 0

0 1 2 3 0 1 2 0 1 2 3 4 5 6 7
Number of Classes Number of Classes Number of Classes

Figure 4. Average richness for all flowering plants plotted against the number of layers in each extreme category.
Randomizations of the data in blue, observed data line green if significantly greater or less than expected by
chance (falling outside ≥97.5%, ≤2.5% of randomizations), red if not significantly outside of random.
 w
ie
Richness
0 - 18
19 - 57
58 - 104
105 - 145
146 - 185
ev
186 - 232
233 - 291
292 - 385
R
386 - 512
513 - 645

Extremeness
In

0
1
2
3
4
5
6
7
8

Figure 5. Grid cell values for North American Compositae diversity (size) and associated extremeness (color).
 A

C
B

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R

D
In

Richness
0 - 512
513 - 1024
1025 - 1536
1537 - 2048
2049 - 2560
2561 - 3072
3073 - 3584
E 3585 - 4096
4097 - 4608
4609 - 5121

Number of
Extreme Classes
0
1
2
3
4
5
6
7
8

Figure S1. Grid cell values for global flowering plant diversity (size) and associated extremeness (color) for (A) North
America, (B) South America, (C) Africa, (D) Eurasia, (E) Australia.
 NORTH AMERICA R-squared: 0.1448007 SOUTH AMERICA R-squared 0.133241

8
7
6

6
# Extreme Classes
# Extreme Classes
5
4

4
3
2

2
1

0
0

20 40 60 80 0 10 20 30 40 50
Latitude Latitude
AFRICA R-squared 0.1357156 EURASIA R-squared 0.1280792
8

w
6
6
# Extreme Classes

# Extreme Classes

ie 5
4
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4

3
2
R
2

1
In
0

0 10 20 30 0 20 40 60 80
Latitude Latitude
AUSTRALIA R-squared 0.006231861
8
# Extreme Classes
6
4
2
0

10 15 20 25 30 35 40
Latitude

Figure S2. Regression and R2 values for flowering plant richness vs number of extreme classes for each continent.
 ALL FLOWERING PLANTS North American
A GLOBAL NORTH AMERICA SOUTH AMERICA AFRICA EURASIA AUSTRALIA
Compositae

4500 5000 1500

3500 4500
CLASSES vs RAW DIVERSITY

4000 4500
1500 4000
3000 500
3500 4000
3500
3000 3500 2500
3000
Richness

2500 3000 2000 1000

2500
2000 2500
1500 2000
1500 2000
1000 1500
1500 1000
1000 1000
1000 500 500
500 500
500
0 0 0 0

B
0 1 2 3 4 5 6 7 8 0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7 8 0 1 2 3 4 5 6 7 8 0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7 8 0 1 2 3 4 5 6 7 8
Number of Extreme Classes
2000 2000 2000 2000 2000 2000
CLASSES vs DIVERSITY RESIDUALS

500
1500 1500 1500 1500 1500 1500

1000 1000 1000 1000 1000 1000

Richness

w
500 500 500 500 500 500

0 0
0 0 0 0

500 0
500 500 500 500 500

ie
1000 0 1 2 3 4 5 6 7 8 1000 0 1 2 3 4 5 6 7 1000 0 1 2 3 4 5 6 7 8 1000 0 1 2 3 4 5 6 7 8 1000 0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7 8 0 1 2 3 4 5 6 7 8
Number of Extreme Classes

ev
3000 4500 5000 1500
LAYERS VS RAW DIVERSITY

3500 4500
4000 4500 400
2500 4000
3500 4000 3000
3500
2000 3000 3500 2500 300
3000
Richness

2500 3000 1000

2000 2500
1500 2500

1000
2000
1500
1000
2000
1500
1000
R 1500

1000
2000
1500
1000
200

100

500 500 500 500 500

500
0 0 0 0
0 1 2 3 4 5 6 7 8 9 10111213 0 1 2 3 4 5 6 7 8 9 10111213 0 1 2 3 4 5 6 7 8 9 10111213 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 10 11 12 0 1 2 3 4 5 6 7 8 9 10 0 1 2 3 4 5 6 7 8 9 10111213
D
In
Number of Extreme Layers

3500
LAYERS VS DIVERSITY RESIDUALS

2500 3000
3000 2500 3000
2000 2500 500
2500 2000 1500 2500
2000
2000 1500 2000 500
1000 1500
1500
Richness

1000 500 1500 1000

1000
500 0 1000 500
500
0 500 500 0
0 0
1000 0 500
500 500 0
1500 500 1000
1000 1000
2000 1500
15000 1 2 3 4 5 6 7 8 9 10111213 0 1 2 3 4 5 6 7 8 9 10111213 0 1 2 3 4 5 6 7 8 9 10111213 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 10 11 12 0 1 2 3 4 5 6 7 8 9 10 0 1 2 3 4 5 6 7 8 9 10111213
Number of Extreme Layers

Figure S3. (A) Average richness for all flowering plants plotted against the number of extreme classes globally, for each continent independently, and for North
American Compositae only; (B) with richness corrected for latitude (residuals of richness) plotted against extreme classes; (C) for average richness plotted against
number of layers (unassigned to classes); (D) with richness corrected for latitude and plotted against number of layers. Randomizations of the data in blue, observed
data line green if significantly greater or less than expected by chance (falling outside ≥97.5%, ≤2.5% of randomizations), red if not significantly outside of random.
 1500
North American Flowering Plant
Richness Including Greenland

1000

w
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500

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R
0
In
0 1 2 3 4 5 6 7
Number of Extreme Classes

Figure S4. Response of North American flowering plant richness against number of extreme classes with the inclusion of
glaciated Greenland areas. Randomizations of the data in blue, observed data line green if significantly greater or less than
expected by chance (falling outside ≥97.5%, ≤2.5% of randomizations), red if not significantly outside of random.
 North America
South America
Africa
Eurasia 290
Australia 125

300

ew
200

182
Number of cells

200

i
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R
155
100 71
In
1
1
1 1
13 6
1

3 4 3
2 1 1 2 2
1 1 1

WaterLimitation
UVExposure
SoilPhysical
NutrientLim
HighMoisture
HeatStress
ColdStress
ExtremeVariability
EnergyLim
Number of 4
Extreme Classes 8 7 6 5 3

Figure S5. Frequency plot showing number of grid cells on each continent under each combination of extreme classes for 3-9 overlapping classes. Hash lines mark segments that are not to scale, with adjacent numbers indicating the number of
cells for that continent.
 AFRICA COLD STRESS AFRICA ENERGY LIMITATION
AFRICA EXTREME VARIABILITY
3500 3500 3500

3000 3000 3000

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 0 0
0 1 2 3 0 1 2 3 0 1 2 3 4
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

AFRICA HEAT STRESS AFRICA HIGH MOISTURE AFRICA NUTRIENT LIMITATION

3500 3500 3500

w
3000 3000 3000
Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

2500 2500 2500

2000 2000
ie 2000
ev
1500 1500 1500

1000 1000 1000

R

500 500 500

0 0 0
0 1 2 3 0 1 2 3 0 1 2 3 4 5
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes
In

AFRICA SOIL PHYSICAL STRESS AFRICA UV EXPOSURE AFRICA WATER LIMITATION

3500 3500 3500

3000 3000 3000

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 0 0
0 1 2 3 0 1 2 0 1 2 3 4 5 6
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes
 AUSTRALIA COLD STRESS 1500 AUSTRALIA ENERGY LIMITATION AUSTRALIA EXTREME VARIABILITY
1500 1500
Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

1000 1000 1000

500 500 500

0 1 2 3 0 1 2 3 0 1 2 3
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

1500 AUSTRALIA HEAT STRESS 1500 AUSTRALIA HIGH MOISTURE 1500 AUSTRALIA NUTRIENT LIMITATION

w
Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

1000 1000
ie 1000
ev
R

500 500 500

0 1 2 3 0 1 2 3 0 1 2 3 4
In

Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

1500 AUSTRALIA SOIL PHYSICAL STRESS 1500 AUSTRALIA UV EXPOSURE 1500 AUSTRALIA WATER LIMITATION
Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

1000 1000 1000

500 500 500

0 1 2 3 0 1 0 1 2 3 4 5
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes
 EURASIA COLD STRESS EURASIA ENERGY LIMITATION EURASIA EXTREME VARIABILITY
4500 4500 4500

4000 4000 4000

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3500 3500 3500

3000 3000 3000

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 0 0

0 1 2 3 0 1 2 0 1 2 3
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

EURASIA HEAT STRESS EURASIA HIGH MOISTURE EURASIA NUTRIENT LIMITATION

4500 4500 4500

w
4000 4000 4000
Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3500 3500 ie 3500

3000 3000 3000

2500 2500 2500

ev
2000 2000 2000

1500 1500 1500

1000 1000 1000

R

500 500 500

0 0 0
In

0 1 2 3 0 1 2 3 0 1 2 3 4 5
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

4500 EURASIA SOIL PHYSICAL STRESS EURASIA UV EXPOSURE 4500

EURASIA WATER LIMITATION
4500

4000 4000 4000

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3500 3500 3500

3000 3000 3000

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 0 0

0 1 2 3 0 1 0 1 2 3 4 5 6
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes
 4500 NORTH AMERICA COLD STRESS vs RICHNESS 4500
NORTH AMERICA ENERGY LIMITATION vs RICHNESS 4500 NORTH AMERICA EXTREME VARIABILITY vs RICHNESS

4000 4000 4000

3500 3500 3500

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3000 3000 3000

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 0 0

0 1 2 0 1 0 1 2 3 4
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

4500
NORTH AMERICA HEAT STRESS vs RICHNESS 4500 NORTH AMERICA HIGH MOISTURE vs RICHNESS 4500
NORTH AMERICA NUTRIENT LIMITATION vs RICHNESS

w
4000 4000 4000

3500 3500 3500

ie
Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3000 3000 3000

2500 2500 2500

ev
2000 2000 2000

1500 1500 1500

1000 1000 1000

R

500 500 500

0 0 0
In

0 1 2 3 0 1 2 3 0 1 2 3 4 5
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

4500
NORTH AMERICA SOIL PHYSICAL STRESS vs RICHNESS 4500
NORTH AMERICA UV DAMAGE vs RICHNESS 4500
NORTH AMERICA WATER LIMITATION vs RICHNESS

4000 4000 4000

3500 3500 3500

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3000 3000 3000

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 0 0

0 1 2 3 0 1 2 0 1 2 3 4 5 6
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes
 5000 SOUTH AMERICA COLD STRESS 5000 SOUTH AMERICA ENERGY LIMITATION 5000 SOUTH AMERICA EXTREME VARIABILITY

4500 4500 4500

4000 4000 4000

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3500 3500 3500

3000 3000 3000

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 1 2 3 0 1 2 3 0 1 2 3 4 5
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

5000 SOUTH AMERICA HEAT STRESS 5000 SOUTH AMERICA HIGH MOISTURE 5000 SOUTH AMERICA NUTRIENT LIMITATION

w
4500 4500 4500

4000 4000 4000

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3500 3500
ie 3500

3000 3000 3000

ev
2500 2500 2500

2000 2000 2000

1500 1500 1500

R

1000 1000 1000

500 500 500

In

0 1 2 3 0 1 2 3 0 1 2 3 4 5
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

5000 SOUTH AMERICA SOIL PHYSICAL STRESS 5000 SOUTH AMERICA UV EXPOSURE 5000 SOUTH AMERICA WATER LIMITATION

4500 4500 4500

4000 4000 4000

Flowering Plant Species Richness

Flowering Plant Species Richness

Flowering Plant Species Richness

3500 3500 3500

3000 3000 3000

2500 2500 2500

2000 2000 2000

1500 1500 1500

1000 1000 1000

500 500 500

0 1 2 3 0 1 2 0 1 2 3 4 5 6 7
Number of Extreme Classes Number of Extreme Classes Number of Extreme Classes

Figure S6. Response of flowering plant richness within individual extreme classes for each continent. Randomizations
of the data in blue, observed data line green if significantly greater or less than expected by chance (falling outside
97.5% or 2.5% of randomizations), red if not significantly outside of random.
 Cold Stress

Individual variable layer: Bottom 5th percentile of Annual Mean Temperature Individual variable layer: Bottom 5th percentile of Minimum Temperature of Coldest Month

w
ie
ev
R
Individual variable layer: Top 5th percentile of Temperature Annual Range Individual variable layer: Bottom 5th percentile of Annual Mean Radiation

In
 Cold Stress Class Layer

w
ie
3.0

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2.0

R
In 1.0

0.0
 Energy Limitation

Individual variable layer: Top 5th percentile of Elevation Individual variable layer: Bottom 5th percentile of Annual Mean Temperature

w
ie
ev
R
Individual variable layer: Bottom 5th percentile of Annual Mean Radiation

In
 Energy Limitation Class Layer

w
ie
3.0

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2.0

R
In 1.0

0.0
 Extreme Variability

Individual variable layer: Bottom 5th percentile of Depth to Bedrock Individual variable layer: Top 5th percentile of Coarse Fragments (Volumetric in %)

w
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R
Individual variable layer: Top 5th percentile of Precipitation Seasonality Individual variable layer: Top 5th percentile of Mean Diurnal Range

In
 Extreme Variability

Individual variable layer: Top 5th percentile of Temperature Seasonality Individual variable layer: Top 5th percentile of Temperature Annual Range

w
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R
In
Extreme Variability Class Layer

w
ie
5

ev
4

R
3

2
In 1

0
 Heat Stress

Individual variable layer: Top 5th percentile of Annual Mean Temperature Individual variable layer: Top 5th percentile of Maximum Temperature of Warmest Month

w
ie
ev
R
Individual variable layer: Top 5th percentile of Annual Mean Temperature Individual variable layer: Top 5th percentile of Annual Mean Radiation

In
 Heat Stress Class Layer

w
ie
3.0

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2.5

2.0

R
1.5

In 1.0

0.5

0.0
 High Moisture Stress

Individual variable layer: Top 5th percentile of Topographic Wetness Index Individual variable layer: Top 5th percentile of Annual Precipitation

w
ie
ev
R
Individual variable layer: Top 5th percentile of Precipitation of Wettest Month Individual variable layer: Top 5th percentile of Mean Annual Humidity

In
High Moisture Class Layer

w
ie
4

ev
3

R
2

In 1

0
 Nutrient Limitation

Individual variable layer: Bottom 5th percentile of Depth to Bedrock Individual variable layer: Bottom 5th percentile of Cation Exchange Capacity of Soil

w
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R
Individual variable layer: Top 5th percentile of Coarse Fragments Volumetric (%) Individual variable layer: Bottom 5th percentile of Soil Organic Carbon Content

In
 Nutrient Limitation

Individual variable layer: Top 5th percentile of Soil pH Individual variable layer: Bottom 5th percentile of Soil pH

w
ie
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R
Individual variable layer: Top 5th percentile of Sand Content

In
Nutrient Limitation Class Layer

w
ie
6

ev
5

R
3

In 2

0
 Soil Physical Stress

Individual variable layer: Bottom 5th percentile of Depth To Bedrock Individual variable layer: Top 5th percentile of Bulk Density kg/cubic-meter

w
ie
ev
R
Individual variable layer: Top 5th percentile of Clay Content (0-2 micrometer) mass fraction in % Individual variable layer: Top 5th percentile of Coarse Fragments (volumetric in %)

In
 Soil Physical Stress

Individual variable layer: Top 5th percentile of sand content (50−2000 micro meter) mass fraction in %

w
ie
ev
R
In
Soil Physical Stress Class Layer

w
ie
4

ev
3

R
2

In 1

0
 UV Exposure

Individual variable layer: Top 5% Elevation Individual variable layer: Top 5% of Annual mean radiation

w
ie
ev
R
In
 UV Exposure Class Layer

w
ie
2.0

ev
1.5

R
1.0

In 0.5

0.0
 Water Limitation

Individual variable layer: Bottom 5th percentile of Topographic Wetness Index Individual variable layer: Bottom 5th percentile of Saturated Water Content

w
ie
ev
R
Individual variable layer: Bottom 5th percentile of depth to bedrock up to 200 cm Individual variable layer: Top 5th percentile of bulk density in kg/cubic−meter

In
 Water Limitation

Individual variable layer: Top 5th percentile of coarse fragments volumetric in % Individual variable layer: Top 5th percentile of sand content (mass fraction in %)

w
ie
ev
R
Individual variable layer: Bottom 5th percentile of Annual Precipitation Individual variable layer: Bottom 5th percentile of Precipitation of Driest Month

In
 Water Limitation

Individual variable layer: Top 5th percentile of Mean Annual wind speed Individual variable layer: Bottom 5th percentile of Mean Annual Humidity

w
ie
ev
R
In
 Water Limitation Class Layer

w
ie
7

ev
5

R
3

2
In
1

Figure S7. Individual variable layers and combined class layers for each of the nine extreme classes. Layers were generated by identifying the area representing the top and/or bottom 5% values for
each variable on each continent independently. These were combined to form global individual variable layers. Individual variables within each extreme class were then stacked to form combined
class layers.
Table S1. Variables as they are assigned to the nine extreme classes for their upper and/or
lower 5th percentile.

Class Original Variable ≥95% ≤5%

Cold Stress
Annual Mean Temperature •
Minimum Temperature of Coldest Month •
Temperature Annual Range •
Annual Mean Radiation •
Energy Limitation
Elevation •
Annual Mean Temperature •
Annual Mean Radiation •
Extreme Variability
Depth to Bedrock •
Volume (%) Coarse Fragments •
Precipitation Seasonality •

w
Mean Diurnal Range •
Temperature Seasonality •
Temperature Annual Range •
Heat Stress

ie
Annual Mean Temperature
Max Temperature Warmest Month
Temperature Annual Range
•
•
•
ev
Annual Mean Radiation •
High Moisture Stress
Topographic Wetness •
Annual Precipitation •
R

Precipitation Wettest Month •

Mean Annual Humidity •
Nutrient Limitation
Depth to Bedrock •
Cation Exchange Capacity of Soil •
In

Volume (%) Coarse Fragments •

Soil Organic Carbon Content (%) •
Soil pH •
Soil pH •
Sand Content •
Soil Physical Stress
Depth to Bedrock •
Bulk Density •
Clay Content (%) •
Volume (%) Coarse Fragments •
Sand Content (%) •
UV Damage
Elevation •
Annual Mean Radiation •
Water Limitation
Topographic Wetness Index •
Saturated Water Content (%) •
 Depth to Bedrock •
Bulk Density (%) •
Volume (%) Coarse Fragments •
Sand Content (%) •
Annual Precipitation •
Precipitation of Driest Month •
Mean Annual Windspeed •
Mean Annual Humidity •

w
ie
ev
R
In
Table S2. Number of 1km2 grid cells that represent each number of overlapping extreme layers
and classes globally.

Number of Overlapping Number of 1km2 pixels

Layers
0 75774341
1 45724977
2 29167883
3 23125419
4 14124436
5 7956988
6 4549232
7 2539726
8 1714562
9 1149793

w
10 752640
11 374344
12 144935
13
14
15
52563
8624
440
ie
ev
16 3
17 1
Number of Overlapping Number of 1km2 pixels
Classes
R

0 75774341
1 39931584
2 25060046
In

3 27938989
4 19199792
5 8176240
6 9425099
7 1308821
8 345583
9 412