Weigelt Et Al 2023 - The Contribution of Plant Life and Growth Forms To Global Gradients of Vacular Plant Diversity PDF

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The contribution of plant life and growth forms to global gradients of vascular plant
diversity
Amanda Taylor*1, Patrick Weigelt1,2,3, Pierre Denelle1, Lirong Cai1, Holger Kreft1,2,3
Taylor, A. (*Corresponding author, amanda.taylor@uni-goettingen.de; ORCID ID: 0000-

0002-0420-2203), Weigelt, P. (pweigel@uni-goettingen.de; 0000-0002-2485-3708), Denelle,
P. (pierre.denelle@uni-goettingen.de; 0000-0002-4729-3774), Cai, L. (lcai@uni-
goettingen.de; 0000-0001-9432-2024), Kreft, H. (hkreft@uni-goettingen.de; 0000-0003-4471-
8236).
1
Biodiversity, Macroecology & Biogeography, Faculty of Forest Sciences & Forest Ecology,
University of Göttingen, Büsgenweg 1, 37077 Göttingen, Germany.
2
Centre of Biodiversity and Sustainable Land Use (CBL), University of Göttingen, Büsgenweg
1, 37077 Göttingen, Germany.
3
Campus Institute Data Science, University of Göttingen, Göttingen, Germany
1
bioRxiv preprint doi: https://doi.org/10.1101/2023.03.06.531444; this version posted March 7, 2023. The copyright holder for this preprint
Summary
Plant life and growth forms (shortened to ‘plant forms’) represent key functional strategies of
plants in relation to their environment and provide important insights into the ecological
constraints acting on the distribution of biodiversity. Despite their ecological importance, how
the spectra of plant forms contribute to global gradients of plant diversity is unresolved.
Using a novel dataset comprising >295,000 species, we quantify the contribution of different
plant forms to global gradients of vascular plant diversity. Further, we establish how plant form
distributions in different biogeographical regions are associated with contemporary and
paleoclimate conditions, environmental heterogeneity, and phylogeny.
We find a major shift in representation by woody perennials in tropical latitudes to herb-

dominated floras in temperate and boreal regions, following a sharp latitudinal gradient in plant
form diversity from the tropics to the poles. We also find significant functional differences
between regions, mirroring life and growth form responses to environmental conditions, which
is mostly explained by contemporary climate (18-87%), and phylogeny (6-62%), with
paleoclimate and heterogeneity playing only a minor role (<23%).
This research highlights variation in the importance of different plant forms to diversity
gradients worldwide, providing a much-needed quantification for long-standing ideas and
concepts structuring plant assemblages.
Key words: biodiversity, functional traits, latitudinal diversity gradient, phylogenetic turnover,
plant distribution
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Introduction
Plant life and growth forms (shortened to ‘plant forms’) are non-phylogenetic classifications of
species that share similar trait combinations (e.g., herbs, geophytes, lianas), and represent key
ecological strategies of plants in relation to their environment. While the grouping of taxa into
plant forms has a long-standing history in ecology (Humboldt, 1806; Warming & Knoblauch,
1895; Raunkiaer, 1934; Box, 1981; Ellenberg & Mueller-Dombois, 1982), the concept is still
widely used in modern-day studies on functional diversity (Mayfield et al., 2005; Leroy et al.,
2021), biome and vegetation classifications (Box & Fujiwara, 2015; Mucina, 2019), ecological
restoration (Gómez-Aparicio, 2009), and plant systematics (Govaerts et al., 2021). Despite this
long period of development and being a major facet of plant life-history variation, how the
spectra of plant life and growth forms contribute to global biodiversity patterns remains
unquantified. For instance, while terrestrial herbs are characteristic of temperate grasslands, and
trees of tropical rainforests, exactly how much they contribute to total plant diversity in or
outside of these biomes, and in relation to coexisting plant forms, are questions that have yet to
be addressed on a global scale due to a lack of data that became available only recently.
Indeed, several plant forms may coexist within a single region, yet may differ substantially in
their proportional representation. Trees offer an exceptional example, as while globally diverse
in tropical forests (Cazzolla Gatti et al., 2022), their regional contribution to tropical
biodiversity is comparable to co-occurring terrestrial herbs, epiphytes, and climbers (Spicer et
al., 2020). Similarly, plant forms within distinct biomes may comprise species that originate
from different evolutionary lineages in different biogeographical regions, leading to variation
in their contribution within equivalent biomes worldwide. For instance, the large Bromeliaceae
family endemic to the New World contributes significantly to the epiphyte flora in Neotropical
forests, explaining to some degree the comparatively poorer epiphyte diversity in tropical forest
biomes in other botanical realms (Taylor et al., 2022). Another case study is the surprisingly
poor representation of native stem succulents in Australia, despite the subcontinent sharing
many climatic affinities with the trans-continental succulent biome, which has favoured the
establishment of several invasive succulent species (Ringelberg et al., 2020). Thus, regions that
share similar climatic conditions may not necessarily share the same functional composition,
highlighting the importance of historical biogeography, and the need to consider the legacy of
past climate on contemporary plant diversity patterns.
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Sudden fluctuations in climate can promote species extinctions, particularly of poorer

dispersers that have limited ability to migrate, and whose ranges are less likely to intersect with
climate refugia (Sandel et al., 2011). In other cases, species physiologies may prevent the
evolution of adaptations to tolerate increasingly unstable climate conditions. The low diversity
of epiphytes in temperate forests, for instance, has been partly related to the distribution of ice
cover during the Last Glacial Maximum (Taylor et al., 2022), possibly leading to a reduction
in the ‘epiphyte niche’ (Zotz, 2005), and that epiphytes are likely poorly adapted to frost (but
see Zotz and Hietz, 2001. On the contrary, geophytes, which tend to favour seasonally dry
climates, have been shown to decrease in species richness with increasing historical climate
stability (Sosa & Loera, 2017), despite climate stability being a major driver of biodiversity by
promoting speciation (Fine, 2015). Understanding how past climate changes have influenced
present day plant form distributions can ultimately help to predict future climate-driven shifts
in vegetation structure and functioning.
Evolutionary history may also play an important role in governing the contribution of different
plant forms to biodiversity. Under the same environmental conditions, phylogenetically related
species may be ecologically similar in terms of life history, resource use, interaction partners,
and consequently, functional traits (Burns & Strauss, 2011). On the other hand, distantly related
lineages may independently evolve similar traits in different biogeographical realms, as has
been shown for C4 photosynthesis (Sage et al., 2012), and other specialised metabolisms
(Pichersky & Lewinsohn, 2011), insular woodiness (Zizka et al., 2022), shrubby stature
(Gehrke et al., 2016), and stem succulence (Ringelberg et al., 2020). It is also not uncommon
for members of the same plant form to have drastically different phylogenetic constraints, such
as the strong phylogenetic constraints on tropical evergreen tree lineages that do not have
perennating buds and different wood anatomy compared to deciduous trees (Zanne et al., 2014).
The variable contribution of plant forms to biodiversity patterns could therefore elucidate how
the complex life history dimensions of plants are modified by the environment and provide
important new insights into the evolutionary and ecological constraints underpinning plant trait-
and diversity-environment relationships.
Here, we provide a global assessment of the importance of different plant forms to global
patterns of vascular plant diversity (see Engemann et al., 2016 for the New World), and address
how their relative contributions in different biogeographical regions are associated with
contemporary and paleoclimate conditions, environmental heterogeneity, and phylogeny. We
consider two of the most widely accepted and basic classifications of plant forms - ‘life forms’
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and ‘growth forms’, which for centuries have been used to study vegetation around the world
(Humboldt, 1806; Warming & Knoblauch, 1895; Raunkiaer, 1934; Box, 1981; Ellenberg &
Mueller-Dombois, 1982). The term ‘growth form’ refers to the growth habit of plants,
recognising trees, shrubs, subshrubs, terrestrial herbs, climbers, and epiphytes as distinct
functional types (defined in Table 1). Life forms, in contrast, describe the different biological
tolerances and strategies of plants to withstand unfavourable seasonal environmental conditions
based on the location of perennating buds (Raunkiaer, 1934; Ellenberg & Mueller-Dombois,
1982). Therophytes, hemicryptophytes, geophytes, chamaephytes, and phanerophytes (Table 1)
have vastly different survival strategies, ranging from persisting as seeds (therophytes) and
underground storage organs (geophytes), to having perennating buds at (hemicryptophytes),
near (chamaephytes), or well above the soil surface (phanerophytes). Accordingly, therophytes
and geophytes can persist in arid and cold environments, hemicryptophytes and chamaephytes
in cool-temperate climates, and many phanerophyte species are restricted to regions without
frequent frost or drought (Sosa & Loera, 2017; Howard et al., 2019; Irl et al., 2020; Taylor et
al., 2021). While several more advanced classification schemes exist today (e.g., Halloy, 1990),
we argue that because life and growth forms serve as a baseline classification of plant
assemblages worldwide, they represent some of the most accessible and documented plant traits
to date.
Using a global dataset consisting of over 295,000 vascular plant species, we quantify the
relative contribution of life and growth forms to biodiversity in different regions around the
world. As several studies have shown that plant diversity and the distribution of biomes and
vegetation types can be broadly predicted by gradients of temperature and precipitation
(Whittaker, 1975), climate heterogeneity (Reu et al., 2011), and past climate changes (e.g.,
palms, Blach-Overgaard et al., 2013), we expect that similar climatic variables will be
important predictors of plant form distributions. We broadly expect that (H1) the proportional
representation of woody plants will decrease from the tropics to the poles and be inversely
related to the proportional representation of terrestrial herbs. We also expect that (H2) variation
in the composition and contribution of plant forms in different regions around the world will be
driven primarily by gradients of precipitation, temperature, past climate change, heterogeneity,
and phylogeny, the effects of which will depend on individual plant form adaptations.
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Table 1. Major life and growth forms defined by plant growth habit (growth form) and the
position of perennating buds (life form). Although epiphytes were originally classified as
phanerophytes within the Raunkiaer life form scheme (Raunkiaer, 1934), they were later
separated due to their unique arboreal habit (Ellenberg & Mueller-Dombois, 1982). As their
revised classification is not related to the position of perennating buds, we include them here as
a growth form, rather than a life form.
Plant form Definition
In relation to growth habit (growth form)
tree a perennial woody plant larger than a shrub (> 10 m) and typically with
a single, self-supporting trunk e.g., Fagus sylvatica (Fagaceae)
shrub a perennial woody plant larger than a subshrub (0.50-10 m), smaller than
a tree e.g., Coprosma lucida (Rubiaceae)
subshrub a small shrub < 0.50 m that is mostly, but not always, woody at the base
e.g., Euphorbia erinacea (Euphorbiaceae)
herb a small herbaceous plant that never develops a woody stem e.g., Carex
halleriana (Cyperaceae)
epiphyte grows non-parasitically on other plants, produces aerial roots e.g.,

Angraecum muscicola (Orchidaceae)
climber woody or herbaceous plant that roots in the ground but is structurally
supported by other plants e.g. Epipremnum aureum (Araceae)
In relation to the position of perennating buds (life form)
phanerophyte perennial plant that bears its perennating buds well above the surface of
the ground e.g., Persea indica (Lauraceae)
nanophanerophyte any phanerophyte between 25 cm and 2 m in height; buds above soil

level e.g., Vaccinium myrtillus (Ericaceae)
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chamaephyte subshrub and dwarf-shrub with buds on persistent shoots near the soil
surface, max 25 cm above ground e.g., Acantholippia seriphioides
(Verbenaceae)
geophyte plant that survives unfavourable seasons in the form of underground

storage organs e.g., Hyacinthoides non-scripta (Asparagaceae)
hemicryptophyte plant having its overwintering buds located at or just below the soil
surface e.g., Drosera xerophila (Droseraceae)
therophyte annual plant that rapidly completes its life cycle in favourable conditions
and survives unfavourable seasons in the form of seeds e.g., Silene
villosa (Caryophyllaceae)
Material and Methods
Biodiversity data
We merged two of the world’s largest botanical databases - the World Checklist of Vascular
Plants (WCVP version 2.0, Govaerts et al., 2021) and the Global Inventory of Floras and Traits
(GIFT version 3.0, Weigelt et al., 2020) to obtain a near-complete global checklist of life and
growth form classifications. Both databases provide comprehensive global checklists of over
344,000 accepted species that come from a variety of resources, including published
taxonomies, floras, and regional checklists. Additionally, these databases are now freely
available to the public, with the WCVP being accessible here (https://powo.science.kew.org/
under the tab “DATA”), and GIFT here (https://github.com/BioGeoMacro/GIFT).
We obtained from the WCVP and GIFT databases plant form classifications for 223,338 and
250,599 species, respectively, with most growth forms coming from GIFT, and life forms from
WCVP. Due to epiphytes often being incorrectly classified, we did not derive epiphyte
classifications from either database, but instead from a recently published epiphyte list (Zotz et
al., 2021), which amounted to 24,279 species, totalling 295,755 species once all databases were
merged. All species classified as epiphytes from the epiphyte list were removed as other plant
forms to reduce biases in the representation by non-trees (i.e., an epiphyte cannot
simultaneously be an ‘epiphyte’ and ‘herb’). Thus, all ‘herbs’ only include terrestrial herbs.
However, in other cases where multiple classifications were assigned to a single species, we
took all possible combinations to account for some variability in plant forms exhibited by
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individual species across different environments (i.e., an herb in one region may appear shrubby
in another region). Together, our dataset accounts for 87% of all accepted plant species, with
the missing 23% being due to a lack of plant life or growth form classification, or due to having
a classification outside of our 12 major plant forms, which we did not consider due to small
sample sizes. For example, we did not consider hydrophytes due to consisting of just ~870
species. Species names across all databases and lists follow the WCVP taxonomic backbone.
Merging these databases provided several benefits beyond a comprehensive global dataset of
plant form classifications. For one, species distributions from the WCVP are recorded at the
level of ‘Botanical Country’ of the World Geographical Scheme for Recording Plant
Distributions (WGSRPD, Brummitt et al., 2001). GIFT, however, offers checklists at finer
resolutions in some places, with species distributions also being aggregated at the regional (e.g.,
departments or states) to country-level (e.g., Peru) in other places. To obtain the highest
resolution of plant form distributions, we always opted to use checklists from the smaller
regions provided that 1) a complete checklist of native vascular plants was available, and 2) the
smaller regions were perfectly nested within larger regions, where if not we would take the
larger geographical entity (e.g., if we were missing two departments of Ecuador, we would not
use the remaining departments, but instead aggregate species to the entire country of Ecuador).
To this end, 56% of the regions included in this study come exclusively from GIFT, and 64%
from WCVP and GIFT combined, out of a total of 564 continental mainland and island regions
(Table S3). We excluded oceanic islands because of their insularity and unique biogeographic
history, which can lead to deviations in plant functional traits (e.g., Taylor et al., 2019).
Because some plant species have more than one life or growth form assignment, we aggregated
the number of species belonging to each plant form to obtain total ‘growth form’ and ‘life form’
richness, which was then used to calculate the relative proportion of each plant form within a
region. More specifically, while ‘absolute species richness’ is the sum of all species within a
region, ‘growth form richness’ and ‘life form richness’ is the sum of all species of a particular
growth or life form within a region, respectively. Each plant form was then divided by its
respective total richness to obtain its relative contribution to biodiversity within each region
(e.g., number of trees/sum of (trees + shrubs + subshrubs + herbs + climbers + epiphytes)). To
determine how the contribution and composition of plant forms varies among large
biogeographical regions, we followed the same procedure for each major botanical continent;
Africa, Australia, North America, South America, Asia Temperate, and Asia Tropical, as well
as the dominant biome (in terms of total area of occupancy) of each geographical entity (e.g.,
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Namibia = “Deserts & Xeric Shrublands”, Cyprus = “Mediterranean Forests, Woodlands &
Scrub”). We excluded from this analysis biomes that were dominant in only one region to
reduce region-specific biases. As such, we removed the ‘mangrove’ and ‘rock and ice’ biomes
due to them being dominant in single regions, Guinea-Bissau, South and Greenland,
respectively.
To gain a general overview of the diversity of plant forms around the world, we calculated the
Shannon diversity index (H’), which considers both the number of species within each plant
form in a given region and their relative abundance (measured here as proportional
representation). We then took the exponential of this index to quantify the Shannon effective
number of plant forms, which measures the number of equally abundant/common plant forms
needed to produce a particular observed value of H’ (Jost, 2006).
Phylogenetic data
To account for phylogenetic relatedness among species, and to tease apart the evolutionary and
ecological contributions of plant forms to global richness patterns, we calculated phylogenetic
eigenvectors at the region level and included them as additional predictor variables in our global
models. We used the extended version of the Smith and Brown (2018) phylogeny, implemented
in the R package ‘V.PhyloMaker2’ (Jin & Qian, 2022), choosing the World Plants
GBOTB.extended.WP.tre extension due to our taxonomic nomenclature being most similar to
this database (https://www.worldplants.de, Hassler, 2000). We then bound missing species to a
respective congener before executing the phylo.maker function to make a phylogenetic tree
hypothesis under scenario 3 (Jin & Qian, 2022). Although scenario 3 adds missing species as
polytomies, several studies have shown the robustness of this approach when used in large-
scale, biogeographical analyses (Qian & Jin, 2016, 2021; Cai et al., 2022).
In a second step, we calculated phylogenetic turnover between regions separately for each life
and growth form, opting for the Simpson index given its insensitivity to variation in species
richness among sites (Baselga, 2010). However, similar in the way that phylogenetic richness
is strongly influenced by species richness, species compositional turnover should also have a
profound influence on phylogenetic turnover. We therefore quantified the standardised effect
size of phylogenetic turnover (Phyloses) for each plant form, correcting for variation in
compositional turnover among regions using the ‘phyloregion’ package in R (Daru et al., 2020).
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This was done by shuffling the tip names of each plant form-subsetted phylogeny 1,000 times
to create multiple null assemblages and taking the difference between the observed
phylogenetic turnover (Phyloobs) and the mean null phylogenetic turnover (Phylomean_null),
divided by the standard deviation (Phylosd); Phyloses = (Phyloobs - Phylomean_null) / Phylosd.
Finally, we ran a Principal Coordinates Analysis (PCoA) on the resulting Phyloses distance
matrix to create phylogenetic eigenvectors at the regional level, which represents the total
variation in phylogenetic composition among regions. Used as explanatory variables, these
eigenvectors can be used to infer how phylogenetic relatedness of species across regions
influences the contribution of plant forms to richness patterns within a given region. We
included in our models the first 4 PCoA axes, as together these could explain > 90% of the
variation in phylogenetic relatedness among regions.
Environmental predictor variables
We initially selected 15 candidate predictor variables representing contemporary climate,

paleoclimate, and environmental heterogeneity. To assess how life and growth forms are
associated with contemporary climate, we selected three metrics derived from CHELSA V.2.1
(30 arc-seconds, Karger et al., 2017), which in previous studies have been shown to play an
important role in determining plant diversity patterns (Scheiner & Rey-Benayas, 1994; Kreft &
Jetz, 2007; Cai et al., 2022): mean annual temperature (°C), mean annual precipitation (mm),
and temperature seasonality (coefficient of variation of mean annual precipitation). These
predictors should capture life and growth form strategies that may or may not favour seasonal
climates (e.g., temperate zone), regions with extreme temperatures (e.g., deserts, alpine
environments), and regions that experience prolonged periods of drought.
To test for potential legacies of historical climate on present day plant form contributions we
considered three paleoclimate variables: climate change velocity in precipitation and
temperature since the Last Glacial Maximum (m/a, 30 arc-seconds, Hijmans et al., 2005), and
climate stability (Owens & Guralnick, 2019). Rapid past climate changes are likely to favour
plant forms that can swiftly recover following a disturbance (e.g., some herbs, Haukioja &
Koricheva, 2000), while stable climates should favour most plant forms, but especially those
restricted in their climate niche (e.g., epiphytes, Taylor et al., 2022). Finally, climate
heterogeneity measured as the Euclidean distance to a multivariate climate centroid (reprojected
from 50 x 50 km2 grids, Fournier et al., 2020) was included to tease apart plant forms that are
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limited to stable, homogeneous climates (e.g., tropics), from those able to persist in
heterogeneous, non-stable climates (e.g., temperate regions).
Several historical climate variables (during the last interglacial and Pleistocene warming
period) were considered - mean annual temperature, temperature seasonality, mean annual
precipitation, and precipitation seasonality, yet were excluded due to strong correlations
(Pearson cor = > 0.70) with contemporary climate variables. Length of growing season, aridity,
number of frost days, and actual evapotranspiration were also excluded due to similar issues of
collinearity. In the case of correlated variables, we always selected those that were most
strongly related to life and growth form representation. One exception is mean annual
temperature and temperature seasonality, which despite being correlated (Pearson cor = 0.71)
we decided to retain both predictors given that seasonality may strongly reflect plant form
strategies to survive harsh conditions, and that temperature is one of the most important drivers
of plant diversity (Whittaker, 1975). We also found a higher model performance with their
combined inclusion (based on lower AIC values of models with both predictors vs models with
temperature and seasonality included separately). A summary of the retained predictor variables
used in this study and their hypothesised effect on each plant form is provided in supporting
information (Table S1, S2). Further, we attach as an additional supporting file Table S3 our
aggregated data sheet with all response and predictor variables used for the analyses.
Statistical Analysis
Generalised linear mixed effects models (GLMMs) with a binomial error term were used to
determine the ecological conditions favouring each plant form, whilst simultaneously
controlling for phylogenetic non-independence of species within each region. We included all
contemporary climatic and past environmental predictors described above as fixed effects, and
botanical continent as a random effect to account for large-scale geographic variation in
evolutionary history among regions (Gerstner et al., 2014). Relative proportions of plant forms
were included as composite response variables (no. of species in plant form x, number of
species in other plant forms).
To test for issues of spatial autocorrelation and over-dispersion, which commonly lead to the
inflation of type 1 errors in macroecological analyses (Dormann et al., 2007; Bolker et al.,
2009), we ran non-spatial GLMMs as described above, and computed the Moran’s I coefficient
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on our model residuals. As we found only low to moderate spatial dependency, ranging from
Moran’s I = 0.11, p < 0.05 for hemicryptophytes to Moran’s I = 0.27, p < 0.05 for
chamaephytes, we decided not to run spatial models. Conversely, we encountered significant
over-dispersion across all plant form models (chisq/rdf = > 500, p < 0.05), which was resolved
by including an additional observation-level random effect. We considered our final and most
parsimonious models as those with the lowest AICc scores and delta AICc = <2 (Burnham &
Anderson, 2004). Finally, we calculated the relative importance of contemporary climate,
paleoclimate, environmental heterogeneity, and phylogeny on the proportional contribution of
life and growth forms to global biodiversity patterns. All statistical analyses and maps were
made in R version 4.2.1 with major input from the packages ‘lme4’ (Bates, 2010), ‘jtools’
(Long, 2019), ‘MuMin’ (Barton, 2015), ‘V.Phylomaker2’ (Jin & Qian, 2022), ‘phytools’
(Revell, 2012), ‘phyloregion’ (Daru et al., 2020), ‘lwgeom’ (Pebesma, 2018), and ‘ggplot2’
(Wickham & Chang, 2016).
Results
Contribution of life and growth forms to botanical continents and biomes
We found striking differences in the contribution of different plant forms to global biodiversity
patterns and in relation to key global drivers of vegetation. Relative growth form composition
consisted of 16% trees, 28% shrubs, 12% subshrubs, 32% herbs, 7% epiphytes, and 5%
climbers (rounded to the nearest whole number). The composition of life forms consisted of
21% phanerophytes, 20% nanophanerophytes, 15% chamaephytes, 24% hemicryptophytes,
11% geophytes, and 9% therophytes. It is important to remember that some species have >1
plant form classification, and that these proportions are in relation to total growth form richness
(n = 325,538) and life form richness (n = 245,129), not species richness.
Terrestrial herbs (incl. herbs, hemicryptophytes, geophytes, therophytes) were most prevalent
in the northern botanical continents, accounting for up to 77% of all vascular plants in North
America, Europe, and Asia-temperate (Fig. 1a). The contribution by perennial woody plants
(tree, shrub, subshrub), in contrast, peaked in South America, Africa, Asia-tropical, and
Australasia, comprising between 54% and 63% of the vascular flora. Epiphytes and climbers
were generally poorly represented in all regions, except for South America and Asia-tropical,
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where epiphytes characterised 12% and 13% of the total flora, and climbers 6%, and 8%,
respectively.
Comparisons among biomes also revealed variation in the distribution of plant form
composition. Species composition of biomes characterised by extreme cold (‘boreal
forests/taiga’, ‘temperate conifer forests’, ‘tundra’, ‘montane grasslands and shrublands’) was
dominated by herbaceous plant forms, which accounted for up to 85% of the flora. Shrub and
subshrub species followed in second place, comprising between 22% and 36% of the flora in
these cooler regions. Arid and hot biomes (‘deserts and xeric shrublands’, ‘Mediterranean
forests, woodlands and scrub’) were composed mainly of herbs (43-47%), shrubs (27-34%),
and hemicryptophytes (27-30%), but also showed notable contributions from subshrubs (15-
20%), nanophanerophytes (14-16%), chamaephytes (17-22%), geophytes (11-15%), and
therophytes (16-19%).
Changes in the distribution and composition of life and growth forms was evident in the tropical
and subtropical biomes (tropical and subtropical: ‘coniferous forests’, ‘dry broadleaf forests’,
‘grasslands, savannas and shrubland’, ‘moist broadleaf forests’), which favoured perennial
woody vegetation – trees (14-23%) and shrubs (25-29%), as well as phanerophytes (20-32%)
and nanophanerophytes (22-25%), although herbs and hemicryptophytes were also well-
represented (24-34% and 16-20%, respectively). The proportional representation of epiphytes
(12% in moist broadleaf forests) and climbers (8% in dry broadleaf forests) also peaked in these
regions. Finally, ‘temperate broadleaf and mixed forests’ displayed a varied composition of
relatively even proportions of trees, subshrubs, chamaephytes, geophytes, nanophanerophytes,
phanerophytes, and therophytes (9-14%), yet maintained high proportions of herbs, shrubs, and
hemicryptophytes (24-51%, Fig. 1b). Despite variation in the contribution of life and growth
forms to plant diversity among the major botanical continents and biomes, we found a clear
latitudinal gradient in the diversity of plant forms from the tropics to the poles (Fig. 1c).
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Figure 1. The proportional contribution of life and growth forms to (a) botanical continents and
(b) biomes. Figure (c) depicts the global distribution of plant form diversity, quantified here as
the effective number of life and growth forms, taken in this case as the exponential of the
Shannon–Wiener Index, H′. Map follows the Winkel tripel projection (ESRI:54042).
Growth forms
In tropical forests, especially in Southeast Asia and the Neotropics, trees account for the
majority of plant diversity (up to 42%, Fig. 2), with shrubs generally being the second most
prevailing growth form (up to 24%, see Table S3 for all proportions by region). This is in stark
contrast to temperate forests where most plant species are herbs (up to 79% in Europe), while
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trees generally constituted only a small fraction (<3%) in these same regions. Our models
mostly associate this clear variation to gradients of temperature, seasonality, and precipitation,
with trees being strongly negatively related to seasonality (standardised estimate Est. = -0.41,
p = <0.01), and positively related to precipitation (Est. = 0.38, p = <0.01) and temperature (Est.
= 0.35, p = <0.01, Fig. 4a). Herbs and subshrubs displayed an inverse relationship, being
positively related to seasonality, and negatively associated to temperature, with subshrubs also
being negatively associated to precipitation (Table S4). Arid Mediterranean climates and
regions overlaying large desert biomes were composed primarily of shrubs (up to 50%, Fig. 2),
subshrubs (up to 27%), and herbs (up to 75%), which was driven by varying combinations of
temperature, seasonality, precipitation, and phylogeny (Table S4, Fig. 4a).
Despite accounting for only 7% of growth form richness (total number of individuals assigned
as growth forms, not species richness), epiphytes comprised up to 35% of the flora of Ecuador,
even proportionately representing more than their host trees in 16 regions, and exceeding all
other growth forms in two regions (Table S3). The higher contribution of epiphytes to diversity
in tropical forests can be related to mean annual precipitation (Est. = 0.76, p = <0.01), and low
temperature seasonality, given the strong, inverse relationship between epiphytes and this
predictor (Est. = -0.49, p = <0.01). Further corroborating that epiphytes prefer stable climates
was their negative association to past climate change velocity (CCV) in temperature (Est. = -
0.26, p = <0.01), CCV in precipitation (Est. = -0.13, p = 0.01), and positive association to
climate stability (Est. = 0.20, p = <0.01), which only weakly predicted most other growth forms.
Climbing plants, which like epiphytes are structurally dependant on trees, displayed strong,
positive relationships with precipitation (Est. = 0.41, p = <0.01) and temperature (Est. = 0.46,
p = <0.01), and were most prominent in Afrotropical and Neotropical forests (up to 32%).
Climbing plants along with trees decreased in proportional representation with increasing
climate heterogeneity, highlighting their tight association to tropical forests. Finally, our
variable importance analysis revealed phylogeny to be a poor predictor of tree, epiphyte, and
climbing growth forms (6-11%), a moderate to strong predictor of herbs and shrubs (34-38%),
and a strong predictor of subshrub (62%) representation (Fig. 4b). Together, these predictors
explained between 54% and 75% of model variance in our GLMM (marginal R2).
15
Figure 2. Global patterns of the proportional representation of growth forms (a-f), and their
relationship with latitude (panels g-i). Proportional representation was calculated as the number
of species within each growth form relative to the combined richness of all growth forms. Note
that herbs = terrestrial herbs. Maps are projected following the Winkel tripel projection (Winkel
III). See Fig. S1 for equivalent species richness maps.
Life forms
As with growth forms, the distribution of life forms varied considerably among the major
climate gradients spanning tropical, temperate, and arid ecosystems (Fig. 3). Phanerophytes
mirrored the representation of trees and reached their greatest representation in the Neotropics,
Afrotropics, and Southeast Asia (up to 53%). This pattern was best predicted by temperature
16
(Est. = 0.39, p = <0.01, Fig. 4a) followed by precipitation (Est. = 0.35, p = <0.01). The
proportional richness of nanophanerophytes peaked in Mediterranean forests, but also
subtropical dry forests and grasslands (up to 37%, Table S3) and was most strongly related to
temperature (Est. = 0.33, p = <0.01). Chamaephytes peaked in representation in regions
characterised by deserts and shrublands, tropical grasslands and Mediterranean climates (up to
32%), which was driven by a combined effect of precipitation and phylogeny (Fig. 4a, Table
S4). Geophytes, hemicryptophytes, and therophytes displayed opposing patterns of
representation, with geophytes being most prevalent in dry Meditterranean (e.g., Cape
Province, South Africa), hemicryptophytes in boreal regions (up to 71%), and therophytes in
regions with expansive deserts (up to 54%). Despite these geographical differences, the
representation of geophytes, hemicryptophytes, and therophytes was strongly related to
temperature, with geophytes and hemicryptophytes favouring cold temperate climates, and
therophytes favouring hot, seasonal, and dry climates (Fig. 4a, Table S4).
Climate heterogeneity, stability, and past CCV were generally weak predictors of life form
representation, accounting for <15% of total variation explained in our variable importance
analysis, compared to 65-87% explained by temperature, seasonality, and precipitation for all
life forms except chamaephytes for which contemporary climate only explained 17% of
variation. Phylogeny, however, while generally a poor to moderate predictor of life form
representation (10-27%), was the most important predictor of chamaephyte representation
(62%, Fig. 4b). Together, these predictors explained a high degree of model variance, with
marginal R2 values ranging from 39%-86% (Table S4).
17
Figure 3. Global patterns of the proportional representation of life forms (a-f), and their
relationship with latitude (panels g-i). Proportional representation was calculated as the number
of species within each life form relative to the combined richness of all life forms. Note that
herbs = terrestrial herbs. Maps are projected following the Winkel tripel projection (Winkel
III). See Fig. S2 for equivalent species richness maps.
18
Figure 4. Coefficient plots showing the standardised effects of temperature seasonality

(Seasonality, coefficient of variation of temperature), mean annual temperature (Temperature,
°C), mean annual precipitation (Precipitation, mm), climate stability (Stability, no units), past
climate change velocity in precipitation (CCV Prec, m/a) and temperature (CCV Temp, m/a),
climate heterogeneity (Heterogeneity, no units), and phylogeny (PCoA 1-4, as components
from a Principal Coordinates Analysis) on the proportional representation of plant (a) growth
19
forms and (b) life forms. Note that the direction of the PCoA axes is not important. Only
predictors that made it to the most parsimonious models for each plant form are included.
Variable importance (c) plots depict the combined importance of contemporary climate,
paleoclimate, environmental heterogeneity, and phylogeny on plant form contributions to
global gradients of vascular plant diversity. Detailed model results and data used to create these
figures can be found in Table S3, S4.
Discussion
The question of how biodiversity is distributed in space and time is an important and complex
topic in ecology and evolutionary biology and requires an integrative perspective on species
distributions, ecologically relevant traits, and phylogenetic relatedness. While the knowledge
of the distribution of vascular plants has substantially increased with recent advancements in
the availability of global datasets (Enquist et al., 2019; Weigelt et al., 2020; Cai et al., 2022),
studies on the functional component of plant biodiversity are lagging behind. We quantified the
relative contribution of life and growth forms in different regions around the world and found
marked differences in their contributions to global gradients of vascular plant diversity. In
support of H1, we found a major shift in the representation by woody perennials in tropical
latitudes to almost entirely terrestrial herb-dominated floras in temperate and boreal regions.
Notably, while trees may occupy over 80% of biomass in (northern) Europe, America, and
temperate Asia (Woodward et al., 2004), we show that they only account for <3% of the
vascular plant richness. Terrestrial herbs, in contrast, constitute up to 90% of the vascular flora
in these same regions.
Several mechanisms may enable herbaceous species to withstand extreme climate variations.
(Zanne et al., 2014). Geophytes, for instance, can survive environmental pressures by utilising
belowground storage organs and keeping their perennating buds below the soil surface
(Raunkiaer, 1934). In cold climates with frequent frost, hemicryptophytes may shed
aboveground biomass, thus insulating their perennating buds located at or just beneath the soil
surface (Raunkiaer, 1934; Lubbe et al., 2021). Finally, short-lived therophytes can persist as
seeds that lay dormant in seed banks until favourable conditions arise for germination and
regrowth (Cross et al., 2015). Although woody perennials typically lack these adaptations, our
study revealed that shrubs and subshrubs frequently coexist with herbs in high proportions,
20
suggesting the presence of additional strategies that enable these plant forms to thrive in
extreme climates.
Shrubs, for example, are composed of numerous stems that may favour their persistence in
unfavourable climates, as injury to one or multiple stems is less likely to result in plant mortality
(Götmark et al., 2016). Likewise, the sizable allocation of belowground organs of subshrubs
allows individuals to resprout and recover following major disturbances (Giroldo et al., 2017),
which could explain their high occurrence in regions characterized by fire and drought (e.g.,
Australia). Sclerophyllous leaves or succulent stems may provide additional protection to
shrubby growth forms in hot, arid environments through the reduction of water loss and
increased water storage (Guerra & Scremin-Dias, 2018). These special adaptations to withstand
climate extremes make shrubs and subshrubs important contributors of biodiversity in
challenging environments where they likely also provide a variety of ecological functions (e.g.,
habitat, water regulation, soil stability, McKell, 1975).
Trees, on the other hand, reached their greatest proportional representation in tropical forests,
which is a well-known pattern from several empirical studies (Keil & Chase, 2019; Cazzolla
Gatti et al., 2022). Nevertheless, our findings indicate that the proportion of trees in tropical
forests does not match the prevalence of herbs in temperate regions when viewed from a global
perspective, corroborating previous studies that a significant component of biodiversity in
tropical forests actually comes from non-trees (Gentry & Dodson, 1987; Spicer et al., 2020).
Indeed, tropical forests are multi-tiered with epiphytes, climbers, terrestrial herbs, and shrubs
also forming important functional components. Epiphytes, for example, constitute just 9.6% of
the world’s vascular flora (Zotz et al., 2021), yet can represent up to 39% of regional
Neotropical diversity (Taylor et al., 2022), and 50% of local diversity (Kelly et al., 2004). The
increase in proportional representation of epiphytes with decreasing scale highlights the
importance of scale dependencies in biodiversity analyses, not only due to variation in the
contribution by different species, but also the mechanisms underlying these relationships
(Crawley & Harral, 2001). Despite the large scale of our analysis, we found in 16 departments
spanning Ecuador, Colombia, and Bolivia, epiphytes proportionately exceeded that of their host
trees, and in two regions exceeded that of all other growth forms. Our analysis suggests that by
neglecting the contribution by non-trees, we are failing to predict half or more of all plant
biodiversity in tropical forests.
21
Although the relative contribution by different plant forms is strongly associated with climate,
we show that evolutionary history also plays an important role. The strong effect of
phylogenetic relatedness on the representation of subshrubs and chamaephytes may be
explained by few plant lineages having the capacity to diversify into the harsh environments
typically occupied by these plant forms. Significant phylogenetic conservatism has been found
in previous studies on shrubs and subshrubs in arid environments (Zheng et al., 2019),
complementing our results that subshrubs are composed of closely related species. Phylogeny
may also be important in regions where plant forms are composed of only a few, species-rich
genera. One example is of the genus Aeonium (Crassulaceae), most of which are succulent
chamaephytes endemic to the Canary Islands that have diversified into arid, steep, and rocky
habitats inaccessible to most other plant lineages, making them the most successful and
specious plant genus across the archipelago (Jorgensen & Olesen, 2001). While we did not
include oceanic islands in this study, future investigations on which plant forms are most prone
to diversifying on islands may bring us closer to understanding why some plant lineages have
more species than others. Conversely, phylogeny had only a weak effect on the proportional
representation of epiphytes, despite ~70% of all epiphytes being orchids (Zotz et al., 2021).
One possible reason for this is that epiphytes are so strongly coupled to atmospheric conditions,
that this may dilute any phylogenetic signal. Our results support this theory as in most cases,
epiphytes displayed stronger relationships with climate than any other plant form (Fig. 4a).
Past climate change left only a small legacy on present day plant form distributions,
corroborating our H2 that life and growth form contributions are primarily explained by
contemporary climate. Terrestrial herbs, for example, were the only plant form positively
associated with rapid changes in temperature, demonstrating a major advantage of herbs to
exploit disturbed habitats and persist through major climate changes. Global shifts in rainfall
regimes are already changing the proportional representation of plant forms in some biomes
such as tropical savannahs (Zhang et al., 2019), which is not reflected in species richness alone.
This reaffirms the importance of studying plant forms alongside species richness to fully
understand the distribution of biodiversity.
Conclusion
22
This study highlights variation in the importance of different plant forms to diversity gradients
worldwide, providing a much-needed quantification for long-standing ideas and concepts
structuring plant assemblages. In line with our expectations, we confirm that the relative
contribution of plant forms to global gradients of vascular plant diversity is strongly related to
life and growth form dependent responses of plants to regional environmental conditions.
Temperature and precipitation continue to be key predictors of plant biodiversity, although we
also show that phylogeny matters, particularly for plant forms occupying niche habitats or harsh
climates (e.g., subshrubs, chamaephytes). Finally, we highlight the usefulness of considering
life and growth forms in biodiversity analyses as a standardised comparison of floras among
regions, which in the future may shed light on the evolutionary pressures constraining plant
trait distributions or increase our understanding of how different vegetation types may respond
to global climate change.
Author contributions
AT, PW, PD, LC, HK conceived the study, AT collated the data from WCVP, PW and HK
created the GIFT database with recent input by PD, AT ran the analyses which were discussed
with PW, AT wrote the draft, and all authors significantly contributed to revisions.
Data availability statement
All aggregated data used for analyses is included in supporting information Table S3.
23
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bioRxiv preprint doi: https://doi.org/10.1101/2023.03.06.531444; this version posted March 7, 2023.

The copyright holder for this preprint

Taylor, A. (*Corresponding author, amanda.taylor@uni-goettingen.de; ORCID ID: 0000-

We find a major shift in representation by woody perennials in tropical latitudes to herb-

Sudden fluctuations in climate can promote species extinctions, particularly of poorer

Plant form Definition

In relation to growth habit (growth form)

epiphyte grows non-parasitically on other plants, produces aerial roots e.g.,

In relation to the position of perennating buds (life form)

nanophanerophyte any phanerophyte between 25 cm and 2 m in height; buds above soil

geophyte plant that survives unfavourable seasons in the form of underground

Material and Methods

Environmental predictor variables

We initially selected 15 candidate predictor variables representing contemporary climate,

Contribution of life and growth forms to botanical continents and biomes

Figure 4. Coefficient plots showing the standardised effects of temperature seasonality

Data availability statement

Barton, K. 2015. Package ‘mumin’.

Leroy, C, Maes, AQ, Louisanna, E, Schimann, H, Séjalon-Delmas, N. 2021. Taxonomic,

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