DOI: 10.5061/dryad.b5mkkwhj9
title: Functional traits—not nativeness—shape the effects of large mammalian herbivores on plant communities

description: Large mammalian herbivores (megafauna) have experienced extinctions and
 declines since prehistory. Introduced megafauna have partly counteracted
 these losses yet are thought to have unusually negative effects compared
 to native megafauna. Using a meta-analysis of 3,995 plot-scale plant
 abundance and diversity responses from 221 studies, we found no evidence
 that megafauna impacts were shaped by nativeness, ‘invasiveness’,
 ‘feralness’, coevolutionary history, or functional and phylogenetic
 novelty. Nor was there evidence that introduced megafauna facilitate
 introduced plants more than native megafauna. Instead, we found strong
 evidence that functional traits shaped megafauna impacts, with
 larger-bodied and bulk-feeding megafauna promoting plant diversity. Our
 work suggests that trait-based ecology provides better insight into
 interactions between megafauna and plants than concepts of nativeness.
description: Literature screening and
 digitization This meta-analysis
 was part of a larger effort to understand megafauna impacts on multiple
 facets of ecosystems (e.g. including soil nutrients, invertebrates, etc).
 This ensured that the dataset included plant responses that were also
 measured in studies focused on other response variables (e.g., spider
 diversity). We searched Web of Science with a string of search terms that
 included the common names and Latin genera of all terrestrial mammalian
 megafauna species (common names from HerbiTraits v1.2 (Lundgren et al.
 2021)) separated with an ‘OR’ operand, along with the following search
 terms: “disturb*, graz*, brows*, impact*, effect, affect, disrupt,
 facilitate, invasi*, ecosystem*, vegetat*, plant*, fauna*, reptil*,
 amphib*, bird*, rodent*, fish*, invertebrat*, insect*, soil*, carbon,
 climate, albedo, river*, riparian, desert*, forest*, tundra,
 decomposition, grassland*, savanna*, chaparral, scrub, shrub, diversity,
 heterogeneity, extinction, richness, environment, reptile*, ecolog*,
 hydrolog*, disturbance, density, biodiversity, response*, ecosystem,
 herbaceous, canopy, germination, cover, pollinator*, tree, nutrient*,
 understorey, erosion, grass*, vegetation, community, exclosure,
 competition, effect*, abundance, productivity”. To reduce unrelated
 results we also included a Web of Science category filter (“WC”) of
 “ECOLOGY OR ZOOLOGY OR ENVIRONMENTAL SCIENCES OR BIODIVERSITY CONSERVATION
 OR EVOLUTIONARY BIOLOGY OR GEOGRAPHY PHYSICAL OR REMOTE SENSING OR PLANT
 SCIENCES OR MULTIDISCIPLINARY SCIENCE OR FORESTRY OR ENTOMOLOGY OR MARINE
 & FRESHWATER BIOLOGY OR MYCOLOGY OR BIOLOGY OR OCEANOGRAPHY OR
 ORNITHOLOGY OR BEHAVIORAL SCIENCES OR FISHERIES”.  The Web of Science review was concluded on the 18th
 of February 2021 and returned 60,537 studies. We removed duplicate studies
 using the fuzzy matching algorithm with the function ‘find_duplicates’ in
 the R package ‘revtools’ (version 0.4.1) (Westgate 2019). After removing
 duplicates, our final search returned 46,825 studies. Title screening
 reduced the number of studies to 2,369. We screened the full text of these studies to only
 include studies focused on wild megafauna (≥45 kg) and that compared areas
 with low versus high megafauna densities due to exclosures, policy-driven
 differences (hunting versus no-hunting in adjacent properties), and
 differences in introduction or eradication histories (adjacent islands
 with and without megafauna). Some studies compared areas with and without
 focal megafauna populations for unknown reasons (e.g., a site with and
 without horses with no indication of why horses might be absent (Robertson
 et al. 2019)), which were excluded due to low confidence in the ultimate
 drivers of observed differences. We excluded all before-after comparisons
 (e.g., a plot measured prior to exclosure construction and then
 afterwards) because of the high rates of change in many systems through
 time (via afforestation, shifts in climate, succession, etc.). Studies
 that excluded megafauna but also all vertebrates were excluded. Two
 additional studies reported data from extremely limiting resources (i.e.,
 wetlands in deserts). These were excluded given that such scenarios should
 be analyzed separately, for which we did not have sufficient sample size.
 Studies that evaluated the effects of megafauna on transplants or
 agricultural crops (including plantations) were not digitized. Studies
 that included an appropriate comparison and reported a central tendency
 (mean or median), a measurement of error (standard deviation, standard
 error, variance, etc), and sample size were digitized (n=154). 
 This literature list was supplemented by the
 literature contained in other relevant meta-analyses (Daskin and Pringle
 2016, Eldridge et al. 2020) and those encountered in the bibliographies of
 the studies we digitized. Given the limited number of studies from oceanic
 islands and regarding widely distributed introduced species (feral pigs,
 goats) in our initial Web of Science search, we conducted focused Google
 Scholar searches on July 15th, 2022 with the following terms: “ungulate
 impacts island*”, “introduced goat impact island*”, “introduced deer
 impact*”, “feral camel impact*”, “wild OR feral boar OR hog OR swine
 impact*”, “feral cattle impact*”, “invasive ungulate hawaii OR guam OR new
 zealand OR pacific island OR new caledonia OR galapagos OR caribbean OR
 oceanic island” and a Web of Science search on the 22nd of December 2022
 using the search string “herbivore* AND plant* AND response*”. This
 uncovered an additional 482 studies of which 66 studies were fit for
 inclusion, leading to a total of 221 studies in our final
 dataset. We digitized central
 tendencies (mean, median), error (standard deviation, standard error,
 interquartile ranges), and sample sizes for each response (diversity,
 richness, and abundance) in each study. We used ImageJ to extract data
 from figures  (Schneider et al. 2012). Interquartile ranges and medians
 (e.g., as extracted from boxplots) were converted to means and standard
 deviation using the function qe.mean.sd in the package ‘estmeansd’ version
 1.0.0 (McGrath et al. 2022). Means and SD/SE were reported by 213 studies
 (3,846 observations) while 11 studies (149 observations) reported medians
 and interquartile ranges. We also
 digitized relevant covariates from the text, which included time since
 treatment (e.g., exclosure construction, introduction, eradication, etc),
 study coordinates (latitude, longitude), megafauna density (standardized
 to kg per hectare), relative abundance of megafauna (in the case of
 multispecies megafauna communities and if density was not provided), and
 the scale of measurement (treated both as area, m<sup>2</sup>,
 and maximum measurement length, m, to allow the comparison of transects to
 plots).  If study coordinates were
 not exactly provided, we extracted latitude and longitude from the
 approximate center of each study location in Google Maps. Maximum
 measurement length was calculated as either the hypotenuse of
 square/rectangular plots, the length of transects, or the diameter of
 circular plots. Distributions of megafauna traits, environmental variables
 (see below), and methodological variables were similar between native and
 introduced megafauna communities in our final dataset. We treated measurements of species richness and
 species diversity (e.g., Shannon Weiner index) as ‘diversity’ responses
 and density estimates (individual plants per plot), % cover, and biomass
 as measurements of abundance. Analyzing these responses alone led to
 similar results. We excluded seed abundance and diversity responses, given
 that seedbanks can be at disequilibrium from realized plant communities.
 We included all true plant species, excluding multicellular algae and
 lichen.  Effect sizes
 Given the presence of negative values and
 zeros in our dataset, we calculated effect sizes using Hedges’ g, a
 unitless measure of standardized mean difference between groups. Each
 effect size was associated with sampling variance calculated from the
 sample size and standard deviation of each observation. Effect sizes and
 sampling variances were calculated with the function ‘escalc’ in the R
 package ‘metafor’ (version 3.5-12) (Viechtbauer 2010). Megafauna and plant nativeness Megafauna nativeness was based on study author
 designations or IUCN range maps (17), if unreported. While many
 communities had both native and introduced megafauna present, the vast
 majority of studies only manipulated (excluded) the introduced megafauna,
 which was possible because of body size differences or through
 eradication. Only one study manipulated both native and introduced
 megafauna (Ward‐Jones et al. 2019). Given that the majority of megafauna
 biomass in this study consisted of introduced megafauna, we classified
 this study as introduced. Excluding it (only relevant for abundance
 analyses) led to similar results. The evolutionary exposure of study sites
 to megafauna (i.e., oceanic islands versus continents and offshore
 islands) was determined using PHYLACINE v1.2 range maps- (Faurby et al.
 2018). We considered New Zealand, which possessed avian megafauna, an
 oceanic island without coevolutionary history with mammalian megafauna
 (due to distinctive foraging strategies of avian versus mammalian
 herbivores). However, counting New Zealand as an offshore island led to
 similar results.  The nativeness of
 collective plant responses was assigned as reported by the authors (1,864
 observations from 104 studies). In cases where plant nativeness was
 unspecified (2,136 observations from 155 studies) we evaluated nativeness
 based on author-provided flora descriptions of the study site by referring
 to the Plants of the World Online (POWO n.d.) and the study site location.
 If introduced plants were described in the study system, we described the
 study as mixed (and thus excluded it) unless the introduced plants
 collectively constituted &lt;5% relative abundance (cover, biomass,
 density), as reported by authors, in which case we counted these systems
 as ‘native’. From this, we were able to classify an additional 1,718
 observations from 113 studies as native (1,499 observations, 97 studies),
 mixed (218 observations, 15 studies), and introduced (1 observation, 1
 study). A final portion of studies did not provide site flora descriptions
 (418 observations, 42 studies). These studies generally came from large,
 well-protected landscapes (e.g., Kruger National Park, Arctic tundra). We
 treated these responses as native.  The nativeness of individual plant species,
 relevant only to plant abundance responses, was extracted from the Plants
 of the World Online (POWO n.d.), as above. Plant taxonomy was standardized
 with the Taxonomic Name Resolution Service (TNRS) (Boyle et al.
 2013). Coevolutionary history and
 coevolutionary novelty The
 coevolutionary history between megafauna and the biomes to which they have
 been introduced was determined using biome maps from Olson et al. 2001
 (Olson et al. 2001). Introduced megafauna were considered ’coevolved’ with
 the biome if they would have occurred in the absence of human-caused
 extinctions and range contractions (e.g., <em>Equus ferus caballus
 </em>in North America), based on PHYLACINE v1.2’s (Faurby et al.
 2018) megafauna distributions in the absence of extinctions and range
 contractions, or if the megafauna species was native elsewhere within the
 focal biome, as in the case of megafauna introduced to offshore islands
 within their native continent. Species-level coevolutionary history
 between megafauna and individual plant species was determined by comparing
 plant distributions (POWO n.d.) to PHYLACINE range maps  (Faurby et al.
 2018). In cases of multiple introduced megafauna, we based this on the
 dominant megafauna species, with dominance determined by relative
 biomass. Functional and
 phylogenetic novelty were calculated by identifying coevolved megafauna
 communities, in the absence of Late Pleistocene extinctions and range
 contractions, for each study location from PHYLACINE v1.2 range maps
 (Faurby et al. 2018). Functional novelty was calculated as the Gower
 distance to the most functionally similar coevolved megafauna. Gower
 distances were calculated using the function ‘gowdis’ (R package ‘FD’,
 version 1.0-12.1) (Laliberté et al. 2014) from key megafauna functional
 traits that determine their effect on the environment (provided by
 HerbiTraits (Lundgren et al. 2021)). These included body mass (log10
 scale), two ordinal dietary traits (graminoid consumption, browse
 consumption), fermentation type (converted to an ordinal variable
 describing fermentation efficiency), three non-exclusive binary habitat
 use variables (aquatic, terrestrial, arboreal), a categorical variable
 describing limb morphology (plantigrade, digitigrade, unguligrade).
 Variable weightings followed (Lundgren et al. 2020). Phylogenetic novelty
 was defined as the cophenetic distance between the introduced megafauna
 and the most closely related megafauna in the absence of human-caused
 extinctions and range contractions using the function ‘pd’ (R package
 ‘ape’ version 5.6-2, (Paradis and Schliep 2019), with the phylogeny
 provided by PHYLACINE v1.2 (Faurby et al. 2018)).  For both phylogenetic and functional novelty, we
 identified the distances between the introduced megafauna and the most
 similar prehistoric ‘coevolved’ megafauna. This value was relativized by
 the introduced species’ relative biomass in their community and then
 averaged across all introduced megafauna. Relative biomass estimates were
 calculated from relative abundance or absolute density estimates per
 species, which were reported for 78.4% of data
 points. <strong><br></strong> Environmental covariates Environmental covariates were extracted for each
 study location by buffering each study location by 5 km and using the
 function ‘extract’ from the R package ‘terra’ (version 1.7-6) (Hijmans
 2023). Values were averaged across the 5 km buffer. Specifically, we
 extracted values of net primary productivity (Zhao et al. 2005), maximum
 annual temperature and precipitation (Fick and Hijmans 2017), and the
 human footprint index (Venter et al. 2016). The human footprint index was
 available for both 1993 and 2009. We thus used values closest to the year
 the data was collected or, if unreported, the year the study was published
 (n=78 studies). For studies reporting data over multiple years, the year
 was adjusted for the time when the individual response was collected based
 on commencement of study and collection interval. Megafauna community functional traits
 To understand how megafauna functional traits
 influence their effects on plants, we evaluated key megafauna functional
 traits for all species in our dataset. Given that 80 studies (1,433
 observations) manipulated multiple species of megafauna, we analyzed
 megafauna functional trait summaries at a community level. We did this by
 multiplying species trait values by their relative biomass in their
 community (0–1) and then calculating the maximum and mean of these traits
 (henceforth ‘community-weighted’). Mean trait values reflect overall
 community tendencies, while the possibility that ecological outcomes may
 be shaped more by extremes (while accounting for relative biomass) is
 captured by maximum trait values. Traits were extracted from HerbiTraits (Lundgren et
 al. 2021) and included body mass, proportion of megafauna biomass with
 hindgut fermentation (which has distinct effects on ecosystems relative to
 foregut fermentation, (Alexander 1993)), and dietary preference for
 graminoids (grasses and allies). Note that while our meta-analysis focused
 on megafauna ≥45 kg, some studies excluded smaller herbivores as well.
 These herbivores were included in trait summaries if ≥10 kg in mass, for
 which trait data was available. Browse and graminoid (grass and allies) consumption
 are the two primary axes of dietary differentiation in herbivores. Browse
 and graminoid consumption were available from HerbiTraits as two
 non-exclusive variables ranging from 0 (avoided) to 3 (highly preferred).
 To synthesize these variables into a single measure, we relativized each
 species’ graminoid consumption value by multiplying it by their relative
 biomass (0-1) within their community. We then divided this value by the
 sum of relativized browse and graminoid consumption.  This variable was
 used in conjunction with the plant growth form of each response,
 categorized as forb, woody, or graminoid. Species-level plant growth forms
 were derived from the World Checklist of Vascular Plants and from the
 ‘growthform’ package (Zanne et al. 2015).  Muzzle widths were extracted from (Pérez–Barbería
 and Gordon 2001, Mendoza et al. 2002). Species-level muzzle widths were
 absent for 42 species of 114 herbivore species in our dataset (36.8%). We
 used genus-level averages for the 25 of these species for which
 genus-level data was available. A remaining 17 species without genus-level
 estimates were minor members (median 13% relative biomass) of speciose
 herbivore communities in 40 observations (4 studies). For these
 observations, we excluded these particular species, calculating muzzle
 width summaries with the other species present only. Elephants
 (<em>Loxodonta africana </em>and<em> Elephas
 maximus</em>), on the other hand, were important components (by
 biomass) of 154 data points (10 studies). Muzzle width is a poor proxy for
 dietary selectivity in elephants since these animals use their trunk to
 forage and can both be selective and consume large quantities of biomass.
 We thus assigned elephants the same muzzle width as black rhinos (Diceros
 bicornis) because of their similar body mass and diet. The final dataset
 included community-weighted muzzle width estimates for all but 56
 abundance responses (1.7%) and 13 diversity responses (1.7%), which were
 excluded from analysis. Finally, we
 assigned each megafauna into functional groups to evaluate whether
 functional group richness shaped megafauna impacts. Functional groups were
 based on combinations of body mass bins (10–45 kg, 45–100 kg, 100–1,000
 kg, ≥1,000 kg), dietary guilds (browser, mixed feeder, grazer), and
 fermentation type (foregut, hindgut, and simple gut).  References 
 Alejandro, L., R. A. Distel, and S. M. Zalba. 2010. Large
 herbivore grazing and non-native plant invasions in montane grasslands of
 central Argentina. Natural Areas Journal 30:148–155.
 Alexander, R. M. 1993. The relative merits of foregut and
 hindgut fermentation. Journal of zoology 231:391–401.
 Allen, R. B., I. J. Payton, and J. E. Knowlton. 1984. Effects of
 ungulates on structure and species composition in the Urewera forests as
 shown by exclosures. New Zealand journal of ecology:119–130.
 Anderson, T. M., M. E. Ritchie, and S. J. McNaughton. 2007.
 Rainfall and soils modify plant community response to grazing in Serengeti
 National Park. Ecology 88:1191–1201. Anujan, K., J.
 Ratnam, and M. Sankaran. 2022. Chronic browsing by an introduced mammalian
 herbivore in a tropical island alters species composition and functional
 traits of forest understory plant communities. Biotropica
 54:1248–1258. Asnani, K. M. 2003. Regeneration of
 woodland vegetation after deer browsing in Sharon Woods Metro Park,
 Franklin County, Ohio. The Ohio State University.
 Augustine, D. J., and D. A. Frank. 2001. Effects of migratory
 grazers on spatial heterogeneity of soil nitrogen properties in a
 grassland ecosystem. Ecology 82:3149–3162. Augustine,
 D. J., and S. J. Mcnaughton. 2004. Regulation of shrub dynamics by native
 browsing ungulates on East African rangeland. The Journal of applied
 ecology 41:45–58. Augustine, D. J., B. J. Wigley, J.
 Ratnam, S. Kibet, M. Nyangito, and M. Sankaran. 2019. Large herbivores
 maintain a two‐phase herbaceous vegetation mosaic in a semi‐arid savanna.
 Ecology and evolution 9:12779–12788. Bailey, J. K.,
 and T. G. Whitham. 2002. Interactions among fire, aspen, and elk affect
 insect diversity: reversal of a community response. Ecology
 83:1701–1712. Baines, D., R. B. Sage, and M. M.
 Baines. 1994. The implications of red deer grazing to ground vegetation
 and invertebrate communities of Scottish native pinewoods. The Journal of
 applied ecology:776–783. Baker, B. W., H. C.
 Ducharme, D. C. S. Mitchell, T. R. Stanley, and H. R. Peinetti. 2005.
 Interaction of beaver and elk herbivory reduces standing crop of willow.
 Ecological applications: a publication of the Ecological Society of
 America 15:110–118. Bakker, E. S., M. E. Ritchie, H.
 Olff, D. G. Milchunas, and J. M. H. Knops. 2006. Herbivore impact on
 grassland plant diversity depends on habitat productivity and herbivore
 size. Ecology letters 9:780–788. Barrett, M. A., and
 P. Stiling. 2006a. Effects of Key deer herbivory on forest communities in
 the lower Florida Keys. Biological conservation 129:100–108.
 Barrett, M. A., and P. Stiling. 2006b. Key deer impacts on
 hardwood hammocks near urban areas. The Journal of wildlife management
 70:1574–1579. Batzli, G. O., and C. E. Dejaco. 2013.
 White-tailed deer (Odocoileus virginianus) facilitate the development of
 nonnative grasslands in central Illinois. The American midland naturalist
 170:323–334. Baur, L. E., K. A. Schoenecker, and M.
 D. Smith. 2018. Effects of feral horse herds on rangeland plant
 communities across a precipitation gradient. Western North American
 naturalist / Brigham Young University 77:526–539.
 Bayne, P., R. Harden, and I. Davies. 2004. Feral goats (Capra
 hircus L.) in the Macleay River gorge system, north-eastern New South
 Wales, Australia. I. Impacts on soil erosion. Wildlife research
 31:519–525. Beck, H., J. W. Snodgrass, and P.
 Thebpanya. 2013. Long-term exclosure of large terrestrial vertebrates:
 Implications of defaunation for seedling demographics in the Amazon
 rainforest. Biological conservation 163:115–121.
 Beguin, J., S. D. Côté, and M. Vellend. 2022. Large herbivores
 trigger spatiotemporal changes in forest plant diversity. Ecology
 103:e3739. Bellingham, P. J., and C. N. Allan. 2003.
 Forest regeneration and the influences of white-tailed deer (Odocoileus
 virginianus) in cool temperate New Zealand rain forests. Forest ecology
 and management 175:71–86. Bennett, A. 2008. The
 impacts of sambar (Cervus unicolor) in the Yarra Ranges National Park.
 Citeseer. Bernard, M., V. Boulanger, J.-L. Dupouey,
 L. Laurent, P. Montpied, X. Morin, J.-F. Picard, and S. Saïd. 2017. Deer
 browsing promotes Norway spruce at the expense of silver fir in the forest
 regeneration phase. Forest ecology and management 400:269–277.
 Bloodworth, K. J., M. E. Ritchie, and K. J. Komatsu. 2020.
 Effects of white‐tailed deer exclusion on the plant community composition
 of an upland tallgrass prairie ecosystem. Journal of vegetation science:
 official organ of the International Association for Vegetation Science
 31:899–907. Bockett, F. K. 1998. Ungulate effects on
 tawa (Beilschmiedia tawa) forest in Urewera National Park. Department of
 Conservation. Boiko, S., E. Bielinis, Z. Sierota, A.
 Zawadzka, A. Słupska, M. Nasiadko, and J. Borkowski. 2019. Polish pony
 changes lower layer biodiversity in old growth scots pine stands. Forests,
 Trees and Livelihoods 10:417. Boulanger, V., J.
 Dupouey, F. Archaux, V. Badeau, C. Baltzinger, R. Chevalier, E. Corcket,
 Y. Dumas, F. Forgeard, and A. Mårell. 2018. Ungulates increase forest
 plant species richness to the benefit of non‐forest specialists. Global
 change biology 24:e485–e495. Bourg, N. A., W. J.
 McShea, V. Herrmann, C. M. Stewart, and B. Blossey. 2017. Interactive
 effects of deer exclusion and exotic plant removal on deciduous forest
 understory communities. AoB plants 9. Bowers, M. A.
 1993. Influence of herbivorous mammals on an old-field plant community:
 years 1-4 after disturbance. Oikos :129–141. Boyd, C.
 S., K. W. Davies, and G. H. Collins. 2017. Impacts of feral horse use on
 herbaceous riparian vegetation within a sagebrush steppe ecosystem.
 Rangeland Ecology & Management 70:411–417.
 Boyle, B., N. Hopkins, Z. Lu, J. A. Raygoza Garay, D. Mozzherin,
 T. Rees, N. Matasci, M. L. Narro, W. H. Piel, S. J. McKay, S. Lowry, C.
 Freeland, R. K. Peet, and B. J. Enquist. 2013. The taxonomic name
 resolution service: an online tool for automated standardization of plant
 names. BMC bioinformatics 14:16. Bucher, R., J.
 Rochlitz, N. Wegner, A. Heiß, A. Grebe, D. G. Schabo, and N. Farwig. 2021.
 Deer exclusion changes vegetation structure and hunting guilds of spiders,
 but not multitrophic understory biodiversity. Diversity 13:25.
 Buesching, C. D., C. Newman, J. T. Jones, and D. W. Macdonald.
 2011. Testing the effects of deer grazing on two woodland rodents,
 bankvoles and woodmice. Basic and applied ecology 12:207–214.
 Burke, A. 1997. The impact of large herbivores on floral
 composition and vegetation structure in the Naukluft Mountains, Namibia.
 Biodiversity & Conservation 6:1203–1217.
 Burkepile, D. E., D. I. Thompson, R. W. S. Fynn, S. E. Koerner,
 S. Eby, N. Govender, N. Hagenah, N. P. Lemoine, K. J. Matchett, and K. R.
 Wilcox. 2016. Fire frequency drives habitat selection by a diverse
 herbivore guild impacting top–down control of plant communities in an
 African savanna. Oikos 125:1636–1646. Burns, C. E.,
 S. L. Collins, and M. D. Smith. 2009. Plant community response to loss of
 large herbivores: comparing consequences in a South African and a North
 American grassland. Biodiversity and conservation 18:2327–2342.
 Bush, E. R., C. D. Buesching, E. M. Slade, and D. W. Macdonald.
 2012. Woodland recovery after suppression of deer: cascade effects for
 small mammals, wood mice (Apodemus sylvaticus) and bank voles (Myodes
 glareolus). PloS one 7:e31404. Cadenasso, M. L., S.
 T. A. Pickett, and P. J. Morin. 2002. Experimental test of the role of
 mammalian herbivores on old field succession: community structure and
 seedling survival. The journal of the Torrey Botanical
 Society:228–237. Callan, R., N. P. Nibbelink, T. P.
 Rooney, J. E. Wiedenhoeft, and A. P. Wydeven. 2013. Recolonizing wolves
 trigger a trophic cascade in Wisconsin (USA). The Journal of ecology
 101:837–845. Carrera, R., W. B. Ballard, P. R.
 Krausman, J. Devos, M. C. Wallace, S. Cunningham, O. J. Alcumbrac, and S.
 A. Christensen. 2015. Reproduction and nutrition of desert mule deer with
 and without predation. The Southwestern naturalist 60:285–298.
 Casabon, C., and D. Pothier. 2008. Impact of deer browsing on
 plant communities in cutover sites on Anticosti Island. Ecoscience
 15:389–397. Castleberry, S. B., W. M. Ford, K. V.
 Miller, and W. P. Smith. 2000. Influences of herbivory and canopy opening
 size on forest regeneration in a southern bottomland hardwood forest.
 Forest ecology and management 131:57–64. Cecil, E.
 M., M. J. Spasojevic, and J. H. Cushman. 2019. Cascading effects of
 mammalian herbivores on ground‐dwelling arthropods: Variable responses
 across arthropod groups, habitats and years. The Journal of animal ecology
 88:1319–1331. Charles, G. K., L. M. Porensky, C.
 Riginos, K. E. Veblen, and T. P. Young. 2017. Herbivore effects on
 productivity vary by guild: cattle increase mean productivity while
 wildlife reduce variability. Ecological applications: a publication of the
 Ecological Society of America 27:143–155. Chen, J.,
 Q. Wang, M. Li, F. Liu, W. Li, and L. Yin. 2013. Effects of deer
 disturbance on soil respiration in a subtropical floodplain wetland of the
 Yangtze River. European journal of soil biology 56:65–71.
 Chollet, S., C. Baltzinger, M. Maillard, and J.-L. Martin. 2021.
 Deer exclusion unveils abiotic filtering in forest understorey plant
 assemblages. Annals of botany 128:371–381. Chollet,
 S., S. Padié, S. Stockton, S. Allombert, A. J. Gaston, and J. Martin.
 2016. Positive plant and bird diversity response to experimental deer
 population reduction after decades of uncontrolled browsing. Diversity and
 Distributions 22:274–287. Christopher, C. C., and G.
 N. Cameron. 2012. Effects of invasive Amur honeysuckle (Lonicera maackii)
 and white-tailed deer (Odocoileus virginianus) on litter-dwelling
 arthropod communities. The American midland naturalist
 167:256–272. Cocquelet, A., A. Mårell, S. Bonthoux,
 C. Baltzinger, and F. Archaux. 2019. Direct and indirect effects of
 ungulates on forest birds’ nesting failure? An experimental test with
 artificial nests. Forest ecology and management 437:148–155.
 Cole, R. J., and C. M. Litton. 2014. Vegetation response to
 removal of non-native feral pigs from Hawaiian tropical montane wet
 forest. Biological invasions 16:125–140. Cole, R. J.,
 C. M. Litton, M. J. Koontz, and R. K. Loh. 2012. Vegetation Recovery 16
 Years after Feral Pig Removal from a Wet Hawaiian Forest. Biotropica
 44:463–471. Collard, A., L. Lapointe, J.-P. Ouellet,
 M. Crête, A. Lussier, C. Daigle, and S. D. Côté. 2010. Slow responses of
 understory plants of maple-dominated forests to white-tailed deer
 experimental exclusion. Forest ecology and management
 260:649–662. van Coller, H., and F. Siebert. 2020.
 The impact of herbivore exclusion on forb diversity: Comparing species and
 functional responses during a drought. African journal of ecology
 58:236–250. Coughenour, M. B. 1991. Biomass and
 nitrogen responses to grazing of upland steppe on Yellowstone’s northern
 winter range. The Journal of applied ecology:71–82.
 Cushman, J. H., L. E. Saunders, and T. K. Refsland. 2020.
 Long‐term and interactive effects of different mammalian consumers on
 growth, survival, and recruitment of dominant tree species. Ecology and
 evolution 10:8801–8814. Cushman, J. H., T. A.
 Tierney, and J. M. Hinds. 2004. Variable effects of feral pig disturbances
 on native and exotic plants in a California grassland. Ecological
 applications: a publication of the Ecological Society of America
 14:1746–1756. Dalgleish, H. J., and D. C. Hartnett.
 2009. The effects of fire frequency and grazing on tallgrass prairie
 productivity and plant composition are mediated through bud bank
 demography. Plant Ecology 201:411–420. Damhoureyeh,
 S. A., and D. C. Hartnett. 1997. Effects of bison and cattle on growth,
 reproduction, and abundances of five tallgrass prairie forbs. American
 journal of botany 84:1719–1728. Daskin, J. H., and R.
 M. Pringle. 2016. Does primary productivity modulate the indirect effects
 of large herbivores? A global meta-analysis. The Journal of animal ecology
 85:857–868. deCalesta, D. S. 1994. Effect of
 white-tailed deer on songbirds within managed forests in Pennsylvania. The
 Journal of wildlife management:711–718. DeGraaf, R.
 M., W. M. Healy, and R. T. Brooks. 1991. Effects of thinning and deer
 browsing on breeding birds in New England oak woodlands. Forest ecology
 and management 41:179–191. De Stoppelaire, G. H., T.
 W. Gillespie, J. C. Brock, and G. A. Tobin. 2004. Use of remote sensing
 techniques to determine the effects of grazing on vegetation cover and
 dune elevation at Assateague Island National Seashore: impact of horses.
 Environmental management 34:642–649. De Villalobos,
 A. E., S. M. Zalba, and D. V. Peláez. 2011. Pinus halepensis invasion in
 mountain pampean grassland: effects of feral horses grazing on seedling
 establishment. Environmental research 111:953–959.
 DiTommaso, A., S. H. Morris, J. D. Parker, C. L. Cone, and A. A.
 Agrawal. 2014. Deer browsing delays succession by altering aboveground
 vegetation and belowground seed banks. PloS one 9:e91155.
 Donaldson, J. E., C. L. Parr, E. H. Mangena, and S. Archibald.
 2020. Droughts decouple African savanna grazers from their preferred
 forage with consequences for grassland productivity. Ecosystems
 23:689–701. Dunkell, D. O., G. L. Bruland, C. I.
 Evensen, and C. M. Litton. 2011. Runoff, Sediment Transport, and Effects
 of Feral Pig (Sus scrofa) Exclusion in a Forested Hawaiian Watershed1.
 Pacific science 65:175–194. Eldridge, D. J., J. Ding,
 and S. K. Travers. 2020. Feral horse activity reduces environmental
 quality in ecosystems globally. Biological conservation
 241:108367. Eldridge, D. J., J. Ding, and S. K.
 Travers. 2021. Low‐intensity kangaroo grazing has largely benign effects
 on soil health. Ecological Management & Restoration
 22:58–63. Elson, A., and D. C. Hartnett. 2017. Bison
 increase the growth and reproduction of forbs in tallgrass prairie. The
 American midland naturalist 178:245–259. Ens, E. J.,
 C. Daniels, E. Nelson, J. Roy, and P. Dixon. 2016. Creating
 multi-functional landscapes: Using exclusion fences to frame feral
 ungulate management preferences in remote Aboriginal-owned northern
 Australia. Biological conservation 197:235–246.
 Eschtruth, A. K., and J. J. Battles. 2009. Acceleration of
 exotic plant invasion in a forested ecosystem by a generalist herbivore.
 Conservation biology: the journal of the Society for Conservation Biology
 23:388–399. Falk, J. M., N. M. Schmidt, T. R.
 Christensen, and L. Ström. 2015. Large herbivore grazing affects the
 vegetation structure and greenhouse gas balance in a high arctic mire.
 Environmental research letters: ERL [Web site] 10:045001.
 Faurby, S., M. Davis, R. Ø. Pedersen, S. D. Schowanek, A.
 Antonelli, and J.-C. Svenning. 2018. PHYLACINE 1.2: The Phylogenetic Atlas
 of Mammal Macroecology. Ecology 99:2626. Fick, S. E.,
 and R. J. Hijmans. 2017. WorldClim 2: new 1-km spatial resolution climate
 surfaces for global land areas. International Journal of
 Climatology. Flaherty, K. L., J. S. Rentch, W. N.
 Grafton, and J. T. Anderson. 2018. Timing of white-tailed deer browsing
 affects wetland plant communities. Plant Ecology 219:313–324.
 Frank, D. A. 2005. The interactive effects of grazing ungulates
 and aboveground production on grassland diversity. Oecologia
 143:629–634. Frank, D. A., and P. M. Groffman. 1998.
 Ungulate vs. landscape control of soil C and N processes in grasslands of
 Yellowstone National Park. Ecology 79:2229–2241.
 Frank, D. A., R. L. Wallen, E. W. Hamilton III, P. J. White, and
 J. D. Fridley. 2018. Manipulating the system: How large herbivores control
 bottom‐up regulation of grasslands. The Journal of ecology
 106:434–443. Fraser, D., and H. Hristienko. 1983.
 Effects of moose, Alces alces, on aquatic vegetation in Sibley Provincial
 Park, Ontario. Freedman, B., P. M. Catling, and Z.
 Lucas. 2011. Effects of feral horses on vegetation of Sable Island, Nova
 Scotia. Canadian Field-Naturalist 125:200–212.
 Gizicki, Z. S., V. Tamez, A. P. Galanopoulou, P. Avramidis, and
 J. Foufopoulos. 2018. Long-term effects of feral goats (Capra hircus) on
 Mediterranean island communities: results from whole island manipulations.
 Biological invasions 20:1537–1552. Goetsch, C., J.
 Wigg, A. A. Royo, T. Ristau, and W. P. Carson. 2011. Chronic over browsing
 and biodiversity collapse in a forest understory in Pennsylvania: results
 from a 60 year-old deer exclusion plot. The journal of the Torrey
 Botanical Society 138:220–224. Goheen, J. R., T. M.
 Palmer, G. K. Charles, K. M. Helgen, S. N. Kinyua, J. E. Maclean, B. L.
 Turner, H. S. Young, and R. M. Pringle. 2013. Piecewise disassembly of a
 large-herbivore community across a rainfall gradient: the UHURU
 experiment. PloS one 8:e55192. Goheen, J. R., T. M.
 Palmer, F. Keesing, C. Riginos, and T. P. Young. 2010. Large herbivores
 facilitate savanna tree establishment via diverse and indirect pathways.
 The Journal of animal ecology 79:372–382. Guldemond,
 R., and R. Van Aarde. 2007. The impact of elephants on plants and their
 community variables in South Africa’s Maputaland. African journal of
 ecology 45:327. Guldemond, R., and R. Van Aarde.
 2010. The influence of tree canopies and elephants on sub‐canopy
 vegetation in a savannah. African journal of ecology
 48:180–189. Hanberry, P., B. B. Hanberry, S.
 Demarais, B. D. Leopold, and J. Fleeman. 2014. Impact on plant communities
 by white-tailed deer in Mississippi, USA. Plant ecology &
 diversity 7:541–548. Hannaford, J., E. H. Pinn, and
 A. Diaz. 2006. The impact of sika deer grazing on the vegetation and
 infauna of Arne saltmarsh. Marine pollution bulletin 53:56–62.
 Hendricks-Franco, L., S. L. Stephens, and W. P. Sousa. 2021.
 Mammalian herbivory in post-fire chaparral impacts herbaceous composition
 but not N and C cycling. Journal of Plant Ecology 14:213–228.
 Hensel, M. J. S., B. R. Silliman, E. Hensel, and J. E. K.
 Byrnes. 2022. Feral hogs control brackish marsh plant communities over
 time. Ecology 103:e03572. Hijmans, R. J. 2023. terra:
 Spatial Data Analysis. Hoffman, M. T., C. F. Madden,
 K. Erasmus, N. Saayman, and J. C. Botha. 2009. The impact of indigenous
 ungulate herbivory over five years (2004–2008) on the vegetation of the
 Little Karoo, South Africa. African Journal of Range & Forage
 Science 26:169–179. Hood, G. A., and S. E. Bayley.
 2009. A comparison of riparian plant community response to herbivory by
 beavers (Castor canadensis) and ungulates in Canada’s boreal mixed-wood
 forest. Forest ecology and management 258:1979–1989.
 Horsley, S. B., S. L. Stout, and D. S. DeCalesta. 2003.
 White‐tailed deer impact on the vegetation dynamics of a northern hardwood
 forest. Ecological applications: a publication of the Ecological Society
 of America 13:98–118. Hothorn, T., F. Bretz, and P.
 Westfall. 2015. Package multcomp: Simultaneous inference in general
 parametric models. published online in the CRAN repository.
 Hummel, S. L., H. Campa, and S. R. Winterstein. 2018.
 Understanding how a keystone herbivore, white-tailed deer impacts wetland
 vegetation types in southern Michigan. The American midland naturalist
 179:51–67. Husheer, S. W. 2007. Introduced red deer
 reduce tree regeneration in Pureora Forest, central North Island, New
 Zealand. New Zealand journal of ecology:79–87.
 Husheer, S. W., Q. W. Hansen, and S. C. Urlich. 2005. Effects of
 red deer on tree regeneration and growth in Aorangi Forest, Wairarapa. New
 Zealand journal of ecology:271–277. Ibañez-Alvarez,
 M., E. Baraza, E. Serrano, A. Romero-Munar, C. Cardona, J. Bartolome, and
 J. A. Krumins. 2022. Ungulates alter plant cover without consistent effect
 on soil ecosystem functioning. Agriculture, ecosystems &
 environment 326:107796. Ickes, K., S. J. DeWalt, and
 S. Appanah. 2001. Effects of native pigs (Sus scrofa) on woody understorey
 vegetation in a Malaysian lowland rain forest. Journal of tropical ecology
 17:191–206. IUCN. 2018. IUCN Red List.
 https://www.iucnredlist.org/. Jenkins, L. H., M. A.
 Jenkins, C. R. Webster, P. A. Zollner, and J. M. Shields. 2014. Herbaceous
 layer response to 17 years of controlled deer hunting in forested natural
 areas. Biological conservation 175:119–128. Johnson,
 B. E., and J. H. Cushman. 2007. Influence of a large herbivore
 reintroduction on plant invasions and community composition in a
 California grassland. Conservation biology: the journal of the Society for
 Conservation Biology 21:515–526. Kalisz, S., R. B.
 Spigler, and C. C. Horvitz. 2014. In a long-term experimental demography
 study, excluding ungulates reversed invader’s explosive population growth
 rate and restored natives. Proceedings of the National Academy of Sciences
 111:4501–4506. Kilheffer, C. R., H. B. Underwood, L.
 Ries, J. Raphael, and D. J. Leopold. 2019. Effects of white-tailed deer
 (Odocoileus virginianus) exclusion on plant recovery in overwash fans
 after a severe coastal storm. AoB plants 11:lz059.
 Kolstad, A. L., G. Austrheim, B. J. Graae, E. J. Solberg, G. R.
 Strimbeck, and J. D. M. Speed. 2019. Moose effects on soil temperatures,
 tree canopies, and understory vegetation: a path analysis. Ecosphere
 10:e02966. Lagendijk, G., B. R. Page, and R. Slotow.
 2012. Short‐term effects of single species browsing release by
 different‐sized herbivores on sand forest vegetation community, South
 Africa. Biotropica 44:63–72. Laliberté, E., P.
 Legendre, and B. Shipley. 2014. FD: measuring functional diversity from
 multiple traits, and other tools for functional ecology.
 Lamperty, T., K. Zhu, J. R. Poulsen, and A. E. Dunham. 2020.
 Defaunation of large mammals alters understory vegetation and functional
 importance of invertebrates in an Afrotropical forest. Biological
 conservation 241:108329. Lessard, J.-P., W. N.
 Reynolds, W. A. Bunn, M. A. Genung, M. A. Cregger, E. Felker-Quinn, M. N.
 Barrios-Garcia, M. L. Stevenson, R. M. Lawton, and C. B. Brown. 2012.
 Equivalence in the strength of deer herbivory on above and below ground
 communities. Basic and applied ecology 13:59–66.
 Long, R. A., A. Wambua, J. R. Goheen, T. M. Palmer, and R. M.
 Pringle. 2017. Climatic variation modulates the indirect effects of large
 herbivores on small‐mammal habitat use. The Journal of animal ecology
 86:739–748. Lorentzen Kolstad, A., G. Austrheim, E.
 J. Solberg, L. De Vriendt, and J. D. M. Speed. 2018. Pervasive moose
 browsing in boreal forests alters successional trajectories by severely
 suppressing keystone species. Ecosphere 9:e02458.
 Loydi, A. 2019. Effects of grazing exclusion on vegetation and
 seed bank composition in a mesic mountain grassland in Argentina. Plant
 ecology & diversity 12:127–138. Loydi, A., S.
 M. Zalba, and R. A. Distel. 2012. Vegetation change in response to grazing
 exclusion in montane grasslands, Argentina. Plant ecology and evolution
 145:313–322. Lucas, R. W., R. Salguero-Gómez, D. B.
 Cobb, B. G. Waring, F. Anderson, W. J. McShea, and B. B. Casper. 2013.
 White‐tailed deer (Odocoileus virginianus) positively affect the growth of
 mature northern red oak (Quercus rubra) trees. Ecosphere
 4:1–15. Lucas, S. 2010. The Effects of Ungulates on
 Species Composition and Nutrient Cycles in Central NZ Forests.
 Post-graduate Diploma, University of Otago. Lundgren,
 E. J. (n.d.). MetaMegafauna. Lundgren, E. J., D.
 Ramp, O. S. Middleton, E. I. F. Wooster, E. Kusch, M. Balisi, W. J.
 Ripple, C. D. Hasselerharm, J. N. Sanchez, M. Mills, and A. D. Wallach.
 2022. A novel trophic cascade between cougars and feral donkeys shapes
 desert wetlands. The Journal of animal ecology.
 Lundgren, E. J., D. Ramp, J. Rowan, O. Middleton, S. D.
 Schowanek, O. Sanisidro, S. P. Carroll, M. Davis, C. J. Sandom, J.-C.
 Svenning, and A. D. Wallach. 2020. Introduced herbivores restore Late
 Pleistocene ecological functions. Proceedings of the National Academy of
 Sciences of the United States of America 117:7871–7878.
 Lundgren, E. J., S. D. Schowanek, J. Rowan, O. Middleton, R. Ø.
 Pedersen, A. D. Wallach, D. Ramp, M. Davis, C. J. Sandom, and J.-C.
 Svenning. 2021. Functional traits of the world’s late Quaternary
 large-bodied avian and mammalian herbivores. Scientific data
 8:17. Maillard, M., J.-L. Martin, S. Chollet, C.
 Catomeris, L. Simon, and S. J. Grayston. 2021. Belowground effects of deer
 in a temperate forest are time-dependent. Forest ecology and management
 493:119228. Maron, J. L., and D. E. Pearson. 2011.
 Vertebrate predators have minimal cascading effects on plant production or
 seed predation in an intact grassland ecosystem. Ecology letters
 14:661–669. Martin, J.-L., S. A. Stockton, S.
 Allombert, and A. J. Gaston. 2010. Top-down and bottom-up consequences of
 unchecked ungulate browsing on plant and animal diversity in temperate
 forests: lessons from a deer introduction. Biological invasions
 12:353–371. Masunga, G. S., S. R. Moe, and B.
 Pelekekae. 2013. Fire and grazing change herbaceous species composition
 and reduce beta diversity in the Kalahari sand system. Ecosystems
 16:252–268. McCauley, D. J., F. Keesing, T. P. Young,
 B. F. Allan, and R. M. Pringle. 2006. Indirect effects of large herbivores
 on snakes in an African savanna. Ecology 87:2657–2663.
 McGrath, S., X. Zhao, R. Steele, and A. Benedetti. 2022.
 estmeansd: Estimating the Sample Mean and Standard Deviation from Commonly
 Reported Quantiles in Meta-Analysis. McInnes, P. F.,
 R. J. Naiman, J. Pastor, and Y. Cohen. 1992. Effects of moose browsing on
 vegetation and litter of the boreal forest, Isle Royale, Michigan, USA.
 Ecology 73:2059–2075. McMillan, N. A., D. L. Hagan,
 K. E. Kunkel, and D. S. Jachowski. 2020. Assessing Large Herbivore
 Management Strategies in the Northern Great Plains Using Rangeland Health
 Metrics. Natural Areas Journal 40:273–280. Meier, M.,
 D. Stöhr, J. Walde, and E. Tasser. 2017. Influence of ungulates on the
 vegetation composition and diversity of mixed deciduous and coniferous
 mountain forest in Austria. European journal of wildlife research
 63:1–10. Mendoza, M., C. M. Janis, and P. Palmqvist.
 2002. Characterizing complex craniodental patterns related to feeding
 behaviour in ungulates: a multivariate approach. Journal of zoology
 258:223–246. Meyer, J.-Y., T. Laitame, and J.-C.
 Gaertner. 2019. Short-term recovery of native vegetation and threatened
 species after restoration of a remnant forest in a small oceanic island of
 the South Pacific. Plant ecology & diversity 12:75–85.
 Mills, C. H., H. Waudby, G. Finlayson, D. Parker, M. Cameron,
 and M. Letnic. 2020. Grazing by over-abundant native herbivores
 jeopardizes conservation goals in semi-arid reserves. Global Ecology and
 Conservation 24:e01384. Minoshima, M., M. B. Takada,
 N. Agetsuma, and T. Hiura. 2013. Sika deer browsing differentially affects
 web-building spider densities in high and low productivity forest
 understories. Ecoscience 20:55–64. Mitchell, J., W.
 Dorney, R. Mayer, and J. McIlroy. 2007. Ecological impacts of feral pig
 diggings in north Queensland rainforests. Wildlife research
 34:603–608. Mitchell, J. L. 2002. Ecology and
 management of feral pigs (Sus scrofa) in rainforests. PhD Thesis, James
 Cook University. Morrison, J. A. 2017. Effects of
 white-tailed deer and invasive plants on the herb layer of suburban
 forests. AoB plants 9:lx058. Morris, T., and M.
 Letnic. 2017. Removal of an apex predator initiates a trophic cascade that
 extends from herbivores to vegetation and the soil nutrient pool.
 Proceedings of the Royal Society B: Biological Sciences
 284:20170111. Mosbacher, J. B., A. Michelsen, M.
 Stelvig, H. Hjermstad-Sollerud, and N. M. Schmidt. 2019. Muskoxen modify
 plant abundance, phenology, and nitrogen dynamics in a High Arctic fen.
 Ecosystems 22:1095–1107. Munoz, A., R. Bonal, and M.
 Díaz. 2009. Ungulates, rodents, shrubs: interactions in a diverse
 Mediterranean ecosystem. Basic and applied ecology 10:151–160.
 Murphy, S. J., and L. S. Comita. 2021. Large mammalian
 herbivores contribute to conspecific negative density dependence in a
 temperate forest. The Journal of ecology 109:1194–1209.
 Murray, B. D., C. R. Webster, M. A. Jenkins, M. R. Saunders, and
 G. S. Haulton. 2016. Ungulate impacts on herbaceous‐layer plant
 communities in even‐aged and uneven‐aged managed forests. Ecosphere
 7:e01378. Muthoni, F. K., T. A. Groen, A. K.
 Skidmore, and P. van Oel. 2014. Ungulate herbivory overrides rainfall
 impacts on herbaceous regrowth and residual biomass in a key resource
 area. Journal of arid environments 100:9–17. Myster,
 R. W. 2011. Above-ground vs. below-ground interactive effects of mammalian
 herbivory on tallgrass prairie plant and soil characteristics. Journal of
 plant interactions 6:283–290. Nafus, M. G., J. A.
 Savidge, A. A. Yackel Adams, M. T. Christy, and R. N. Reed. 2018. Passive
 restoration following ungulate removal in a highly disturbed tropical wet
 forest devoid of native seed dispersers. Restoration Ecology
 26:331–337. Nakahama, N., K. Uchida, A. Koyama, T.
 Iwasaki, M. Ozeki, and T. Suka. 2020. Construction of deer fences restores
 the diversity of butterflies and bumblebees as well as flowering plants in
 semi-natural grassland. Biodiversity and conservation
 29:2201–2215. Nuzzo, V., A. Dávalos, and B. Blossey.
 2017. Assessing plant community composition fails to capture impacts of
 white-tailed deer on native and invasive plant species. AoB plants
 9. Ogada, D. L., M. E. Gadd, R. S. Ostfeld, T. P.
 Young, and F. Keesing. 2008. Impacts of large herbivorous mammals on bird
 diversity and abundance in an African savanna. Oecologia
 156:387–397. Okullo, P., and S. R. Moe. 2012. Large
 herbivores maintain termite‐caused differences in herbaceous species
 diversity patterns. Ecology 93:2095–2103. Olson, D.
 M., E. Dinerstein, E. D. Wikramanayake, N. D. Burgess, G. V. N. Powell, E.
 C. Underwood, I. Itoua, H. E. Strand, J. C. Morrison, C. J. Loucks, T. F.
 Allnutt, T. H. Ricketts, Y. Kura, J. F. Lamoreux, W. W. Wettengel, P.
 Hedao, and K. R. Kassem. 2001. Terrestrial Ecoregions of the World: A New
 Map of Life on Earth. Bioscience 51:933–938.
 Opperman, J. J., and A. M. Merenlender. 2000. Deer herbivory as
 an ecological constraint to restoration of degraded riparian corridors.
 Restoration Ecology 8:41–47. Page, M. J., J. E.
 McKenzie, P. M. Bossuyt, I. Boutron, T. C. Hoffmann, C. D. Mulrow, L.
 Shamseer, J. M. Tetzlaff, E. A. Akl, S. E. Brennan, R. Chou, J. Glanville,
 J. M. Grimshaw, A. Hróbjartsson, M. M. Lalu, T. Li, E. W. Loder, E.
 Mayo-Wilson, S. McDonald, L. A. McGuinness, L. A. Stewart, J. Thomas, A.
 C. Tricco, V. A. Welch, P. Whiting, and D. Moher. 2021. The PRISMA 2020
 statement: an updated guideline for reporting systematic reviews. Revista
 espanola de cardiologia 74:790–799. Paradis, E., and
 K. Schliep. 2019. ape 5.0: an environment for modern phylogenetics and
 evolutionary analyses in R. Parker, H. A., J. T.
 Larkin, D. Heggenstaller, J. Duchamp, M. C. Tyree, C. S. Rushing, E. J.
 Domoto, and J. L. Larkin. 2020. Evaluating the impacts of white-tailed
 deer (Odocoileus virginianus) browsing on vegetation in fenced and
 unfenced timber harvests. Forest ecology and management
 473:118326. Peebles-Spencer, J. R., D. L. Gorchov,
 and T. O. Crist. 2017. Effects of an invasive shrub, Lonicera maackii, and
 a generalist herbivore, white-tailed deer, on forest floor plant community
 composition. Forest ecology and management 402:204–212.
 Pekin, B. K., M. J. Wisdom, C. G. Parks, B. A. Endress, and B.
 J. Naylor. 2016. Response of native versus exotic plant guilds to cattle
 and elk herbivory in forested rangeland. Applied Vegetation Science
 19:31–39. Pendergast, T. H., IV, S. M. Hanlon, Z. M.
 Long, A. A. Royo, and W. P. Carson. 2016. The legacy of deer
 overabundance: long-term delays in herbaceous understory recovery.
 Canadian journal of forest research. Journal canadien de la recherche
 forestiere 46:362–369. Perea, R., M. Girardello, and
 A. San Miguel. 2014. Big game or big loss? High deer densities are
 threatening woody plant diversity and vegetation dynamics. Biodiversity
 and conservation 23:1303–1318. Perea, R., A.
 López-Sánchez, J. Pallarés, G. G. Gordaliza, I. González-Doncel, L. Gil,
 and J. Rodríguez-Calcerrada. 2020. Tree recruitment in a drought-and
 herbivory-stressed oak-beech forest: Implications for future species
 coexistence. Forest ecology and management 477:118489.
 Pérez–Barbería, F. J., and I. J. Gordon. 2001. Relationships
 between oral morphology and feeding style in the Ungulata: a
 phylogenetically controlled evaluation. Proceedings of the Royal Society
 of London. Series B: Biological Sciences 268:1023–1032.
 Perrin, P. M., D. L. Kelly, and F. J. G. Mitchell. 2006.
 Long-term deer exclusion in yew-wood and oakwood habitats in southwest
 Ireland: natural regeneration and stand dynamics. Forest ecology and
 management 236:356–367. Pisanu, P., P. Bayne, R.
 Harden, and A. Eggert. 2005. Feral goats (Capra hircus L.) in the Macleay
 River gorge system, north-eastern New South Wales, Australia. II. Impacts
 on rainforest vegetation. Wildlife research 32:111–119.
 Porensky, L. M., S. F. Bucher, K. E. Veblen, A. C. Treydte, and
 T. P. Young. 2013. Megaherbivores and cattle alter edge effects around
 ecosystem hotspots in an African savanna. Journal of arid environments
 96:55–63. Post, E., and C. Pedersen. 2008. Opposing
 plant community responses to warming with and without herbivores.
 Proceedings of the National Academy of Sciences
 105:12353–12358. POWO. (n.d.). Plants of the World
 Online. http://www.plantsoftheworldonline.org/.
 Pringle, R. M., T. P. Young, D. I. Rubenstein, and D. J.
 McCauley. 2007. Herbivore-initiated interaction cascades and their
 modulation by productivity in an African savanna. Proceedings of the
 National Academy of Sciences 104:193–197. Questad, E.
 J., A. Uowolo, S. Brooks, R. Fitch, and S. Cordell. 2018. Resource
 Availability, Propagule Supply, and Effect of Nonnative Ungulate
 Herbivores on Senecio madagascariensis Invasion1. Pacific science
 72:69–79. Qvarnemark, L. M., and S. P. Sheldon. 2004.
 Moose grazing decreases aquatic plant diversity. Journal of freshwater
 ecology 19:407–410. Radloff, F. G. T., L. Mucina, and
 D. Snyman. 2014. The impact of native large herbivores and fire on the
 vegetation dynamics in the Cape renosterveld shrublands of South Africa:
 insights from a six‐yr field experiment. Applied Vegetation Science
 17:456–469. Ramirez, A. R., R. B. Pratt, A. L.
 Jacobsen, and S. D. Davis. 2012. Exotic deer diminish post-fire resilience
 of native shrub communities on Santa Catalina Island, southern California.
 Plant Ecology 213:1037–1047. Ramirez, J. I., P. A.
 Jansen, J. den Ouden, L. Moktan, N. Herdoiza, and L. Poorter. 2021.
 Above-and below-ground cascading effects of wild ungulates in temperate
 forests. Ecosystems 24:153–167. Ratajczak, Z., S. L.
 Collins, J. M. Blair, S. E. Koerner, A. M. Louthan, M. D. Smith, J. H.
 Taylor, and J. B. Nippert. 2022. Reintroducing bison results in
 long-running and resilient increases in grassland diversity. Proceedings
 of the National Academy of Sciences 119:e2210433119.
 Rearick, D., L. Kintz, K. L. Burke, and T. S. Ransom. 2011.
 Effects of white-tailed deer on the native earthworm, Eisenoides
 carolinensis, in the southern Appalachian Mountains, USA. Pedobiologia
 54:S173–S180. Rees, J. D., R. T. Kingsford, and M.
 Letnic. 2017. In the absence of an apex predator, irruptive herbivores
 suppress grass seed production: implications for small granivores.
 Biological conservation 213:13–18. Relva, M. A., M.
 A. Nunez, and D. Simberloff. 2010. Introduced deer reduce native plant
 cover and facilitate invasion of non-native tree species: evidence for
 invasional meltdown. Biological invasions 12:303–311.
 Riesch, F., B. Tonn, H. G. Stroh, M. Meißner, N. Balkenhol, and
 J. Isselstein. 2020. Grazing by wild red deer maintains characteristic
 vegetation of semi‐natural open habitats: Evidence from a three‐year
 exclusion experiment. Applied Vegetation Science 23:522–538.
 Risch, A. C., M. Schütz, M. L. Vandegehuchte, W. H. Van Der
 Putten, H. Duyts, U. Raschein, D. J. Gwiazdowicz, M. D. Busse, D. S.
 Page-Dumroese, and S. Zimmermann. 2015. Aboveground vertebrate and
 invertebrate herbivore impact on net N mineralization in subalpine
 grasslands. Ecology 96:3312–3322. Robertson, G., J.
 Wright, D. Brown, K. Yuen, and D. Tongway. 2019. An assessment of feral
 horse impacts on treeless drainage lines in the Australian Alps.
 Ecological Management & Restoration 20:21–30.
 Rogers, G. M. 1991. Kaimanawa feral horses and their
 environmental impacts. New Zealand journal of ecology:49–64.
 Rojas-Sandoval, J., E. J. Meléndez-Ackerman, J. Fumero-Cabán, M.
 García-Bermúdez, J. Sustache, S. Aragón, M. Morales-Vargas, G. Olivieri,
 and D. S. Fernández. 2016. Long-term understory vegetation dynamics and
 responses to ungulate exclusion in the dry forest of Mona
 Island. Rooney, T. P. 2009. High white-tailed deer
 densities benefit graminoids and contribute to biotic homogenization of
 forest ground-layer vegetation. Plant Ecology 202:103–111.
 Rossell, C. R., Jr, B. Gorsira, and S. Patch. 2005. Effects of
 white-tailed deer on vegetation structure and woody seedling composition
 in three forest types on the Piedmont Plateau. Forest ecology and
 management 210:415–424. Roy, A., M. Suchocki, L.
 Gough, and J. R. McLaren. 2020. Above-and belowground responses to
 long-term herbivore exclusion. Arctic, antarctic, and alpine research
 52:109–119. Royo, A. A., R. Collins, M. B. Adams, C.
 Kirschbaum, and W. P. Carson. 2010. Pervasive interactions between
 ungulate browsers and disturbance regimes promote temperate forest
 herbaceous diversity. Ecology 91:93–105. Royo, A. A.,
 D. W. Kramer, K. V. Miller, N. P. Nibbelink, and S. L. Stout. 2017.
 Spatio-temporal variation in foodscapes modifies deer browsing impact on
 vegetation. Landscape ecology 32:2281–2295. Rutina,
 L. P., and S. R. Moe. 2014. Elephant (Loxodonta africana) disturbance to
 riparian woodland: effects on tree-species richness, diversity and
 functional redundancy. Ecosystems 17:1384–1396.
 Sakai, M., Y. Natuhara, A. Imanishi, K. Imai, and M. Kato. 2012.
 Indirect effects of excessive deer browsing through understory vegetation
 on stream insect assemblages. Population Ecology 54:65–74.
 Saltz, D., H. Schmidt, M. Rowen, A. Karnieli, D. Ward, and I.
 Schmidt. 1999. Assessing grazing impacts by remote sensing in hyper-arid
 environments. Rangeland Ecology & Management/Journal of Range
 Management Archives 52:500–507. Sandhage‐Hofmann, A.,
 A. Linstädter, L. Kindermann, S. Angombe, and W. Amelung. 2021.
 Conservation with elevated elephant densities sequesters carbon in soils
 despite losses of woody biomass. Global change biology
 27:4601–4614. Schneider, C. A., W. S. Rasband, and K.
 W. Eliceiri. 2012. NIH Image to ImageJ: 25 years of image analysis. Nature
 methods 9:671–675. Seliskar, D. M. 2003. The response
 of Ammophila breviligulata and Spartina patens (Poaceae) to grazing by
 feral horses on a dynamic mid‐Atlantic barrier island. American journal of
 botany 90:1038–1044. Shibata, E., M. Saito, and M.
 Tanaka. 2008. Deer-proof fence prevents regeneration of Picea jezoensis
 var. hondoensis through seed predation by increased woodmouse populations.
 Journal of Forest Research 13:89–95. Siemann, E., J.
 A. Carrillo, C. A. Gabler, R. Zipp, and W. E. Rogers. 2009. Experimental
 test of the impacts of feral hogs on forest dynamics and processes in the
 southeastern US. Forest ecology and management 258:546–553.
 Singer, F. J. 1995. Effects of grazing by ungulates on upland
 bunchgrass communities of the northern winter range of Yellowstone
 National Park. Northwest science: official publication of the Northwest
 Scientific Association 69:191–203. Singer, F. J., and
 M. K. Harter. 1996. Comparative effects of elk herbivory and 1988 fires on
 northern Yellowstone National Park grasslands. Ecological applications: a
 publication of the Ecological Society of America 6:185–199.
 Singer, F. J., and R. A. Renkin. 1995. Effects of browsing by
 native ungulates on the shrubs in big sagebrush communities in Yellowstone
 National Park. The Great Basin naturalist:201–212.
 Sitters, J., D. M. Kimuyu, T. P. Young, P. Claeys, and H. Olde
 Venterink. 2020. Negative effects of cattle on soil carbon and nutrient
 pools reversed by megaherbivores. Nature Sustainability
 3:360–366. Souza, Y., N. Villar, V. Zipparro, S.
 Nazareth, and M. Galetti. 2022. Large mammalian herbivores modulate plant
 growth form diversity in a tropical rainforest. The Journal of ecology
 110:845–859. Staver, A. C., and W. J. Bond. 2014. Is
 there a “browse trap”? Dynamics of herbivore impacts on trees and grasses
 in an African savanna. The Journal of ecology 102:595–602.
 Sterne, J. A. C., A. J. Sutton, J. P. A. Ioannidis, N. Terrin,
 D. R. Jones, J. Lau, J. Carpenter, G. Rücker, R. M. Harbord, C. H. Schmid,
 J. Tetzlaff, J. J. Deeks, J. Peters, P. Macaskill, G. Schwarzer, S. Duval,
 D. G. Altman, D. Moher, and J. P. T. Higgins. 2011. Recommendations for
 examining and interpreting funnel plot asymmetry in meta-analyses of
 randomised controlled trials. BMJ 343:d4002.
 Stohlgren, T. J., L. D. Schell, and B. Vanden Heuvel. 1999. How
 grazing and soil quality affect native and exotic plant diversity in Rocky
 Mountain grasslands. Ecological applications: a publication of the
 Ecological Society of America 9:45–64. Stuart-Hill,
 G. C. 1992. Effects of elephants and goats on the Kaffrarian succulent
 thicket of the eastern Cape, South Africa. The Journal of applied
 ecology:699–710. Suominen, O., K. Danell, and R.
 Bergström. 1999a. Moose, trees, and ground-living invertebrates: indirect
 interactions in Swedish pine forests. Oikos :215–226.
 Suominen, O., K. Danell, and J. P. Bryant. 1999b. Indirect
 effects of mammalian browsers on vegetation and ground-dwelling insects in
 an Alaskan floodplain. Ecoscience 6:505–510. Swain,
 M. 2021. Indirect impacts of a non-native ungulate browser on soil
 ecosystem function is variable across soil horizons in the boreal forests
 of Newfoundland, Canada. Memorial University of Newfoundland.
 Tabuchi, K., D. T. Quiring, L. E. Flaherty, L. L. Pinault, and
 K. Ozaki. 2011. Bottom-up trophic cascades caused by moose browsing on a
 natural enemy of a galling insect on balsam fir. Basic and applied ecology
 12:523–531. Tanentzap, A. J., D. R. Bazely, S. Koh,
 M. Timciska, E. G. Haggith, T. J. Carleton, and D. A. Coomes. 2011. Seeing
 the forest for the deer: do reductions in deer-disturbance lead to forest
 recovery? Biological conservation 144:376–382.
 Tanentzap, A. J., L. E. Burrows, W. G. Lee, G. Nugent, J. M.
 Maxwell, and D. A. Coomes. 2009. Landscape‐level vegetation recovery from
 herbivory: progress after four decades of invasive red deer control. Wiley
 Online Library. Taylor, D. L., L.-P. Leung, and I. J.
 Gordon. 2011. The impact of feral pigs (Sus scrofa) on an Australian
 lowland tropical rainforest. Wildlife research 38:437–445.
 Tracy, B. F., and D. A. Frank. 1998. Herbivore influence on soil
 microbial biomass and nitrogen mineralization in a northern grassland
 ecosystem: Yellowstone National Park. Oecologia:556–562.
 Treydte, A. C., C. C. Grant, and F. Jeltsch. 2009. Tree size and
 herbivory determine below-canopy grass quality and species composition in
 savannahs. Biodiversity and conservation 18:3989–4002.
 Uno, H., Y. Inatomi, M. Ueno, and H. Iijima. 2019. Effects of
 sika deer (Cervus nippon) and dwarf bamboo (Sasa senanensis) on tree
 seedlings in a cool-temperate mixed forest on Hokkaido Island, Japan.
 European journal of forest research 138:929–938.
 Vandegehuchte, M. L., M. Schütz, F. De Schaetzen, and A. C.
 Risch. 2017a. Mammal‐induced trophic cascades in invertebrate food webs
 are modulated by grazing intensity in subalpine grassland. The Journal of
 animal ecology 86:1434–1446. Vandegehuchte, M. L., W.
 H. Van Der Putten, H. Duyts, M. Schütz, and A. C. Risch. 2017b.
 Aboveground mammal and invertebrate exclusions cause consistent changes in
 soil food webs of two subalpine grassland types, but mechanisms are
 system‐specific. Oikos 126. Van Staalduinen, M. A.,
 H. During, and M. J. A. Werger. 2007. Impact of grazing regime on a
 Mongolian forest steppe. Applied Vegetation Science 10:299–306.
 Varriano, S., L. H. Lefler, K. Patel, C. Kirksey, A. Turner, and
 M. D. Moran. 2020. The complementary relationship of bison grazing and
 arthropod herbivory in structuring a tallgrass prairie community.
 Rangeland Ecology & Management 73:491–500.
 Veblen, K. E., and T. P. Young. 2010. Contrasting effects of
 cattle and wildlife on the vegetation development of a savanna landscape
 mosaic. The Journal of ecology 98:993–1001. Venter,
 O., E. W. Sanderson, A. Magrach, J. R. Allan, J. Beher, K. R. Jones, H. P.
 Possingham, W. F. Laurance, P. Wood, B. M. Fekete, M. A. Levy, and J. E.
 M. Watson. 2016. Global terrestrial Human Footprint maps for 1993 and
 2009. Scientific data 3:160067. Viechtbauer, W. 2010.
 Conducting meta-analyses in R with the metafor package.
 de Villalobos, A. E., and L. Schwerdt. 2018. Feral horses and
 alien plants: effects on the structure and function of the Pampean
 Mountain grasslands (Argentina). Ecoscience 25:49–60.
 de Villalobos, A. E., and L. Schwerdt. 2020. Seasonality of
 feral horse grazing and invasion of Pinus halepensis in grasslands of the
 Austral Pampean Mountains (Argentina): management considerations.
 Biological invasions 22:2941–2955. Vinton, M. A., and
 D. C. Hartnett. 1992. Effects of bison grazing on Andropogon gerardii and
 Panicum virgatum in burned and unburned tallgrass prairie.
 Oecologia:374–382. Vowles, T., F. Lindwall, A.
 Ekblad, M. Bahram, B. R. Furneaux, M. Ryberg, and R. G. Björk. 2018.
 Complex effects of mammalian grazing on extramatrical mycelial biomass in
 the Scandes forest‐tundra ecotone. Ecology and evolution
 8:1019–1030. Ward, D., D. Saltz, M. Rowen, and I.
 Schmidt. 1999. Effects of grazing by re‐introduced Equus hemionus on the
 vegetation in a Negev desert erosion cirque. Journal of vegetation
 science: official organ of the International Association for Vegetation
 Science 10:579–586. Ward‐Jones, J., I. Pulsford, R.
 Thackway, D. Bishwokarma, and D. Freudenberger. 2019. Impacts of feral
 horses and deer on an endangered woodland of Kosciuszko National Park.
 Ecological Management & Restoration 20:37–46.
 Ward, J. S., and S. C. Williams. 2020. Influence of deer hunting
 and residual stand structure on tree regeneration in deciduous forests.
 Wildlife Society bulletin 44:519–530. Webster, C. R.,
 M. A. Jenkins, and J. H. Rock. 2005. Long-term response of spring flora to
 chronic herbivory and deer exclusion in Great Smoky Mountains National
 Park, USA. Biological conservation 125:297–307.
 Weller, S. G., R. J. Cabin, D. H. Lorence, S. Perlman, K. Wood,
 T. Flynn, and A. K. Sakai. 2011. Alien plant invasions, introduced
 ungulates, and alternative states in a mesic forest in Hawaii. Restoration
 Ecology 19:671–680. Weller, S. G., A. K. Sakai, M.
 Clark, D. H. Lorence, T. Flynn, W. Kishida, N. Tangalin, and K. Wood.
 2018. The effects of introduced ungulates on native and alien plant
 species in an island ecosystem: Implications for change in a diverse mesic
 forest in the Hawaiian Islands. Forest ecology and management
 409:518–526. Werner, P. A. 2005. Impact of feral
 water buffalo and fire on growth and survival of mature savanna trees: An
 experimental field study in Kakadu National Park, northern Australia.
 Austral ecology 30:625–647. Werner, P. A., I. D.
 Cowie, and J. S. Cusack. 2006. Juvenile tree growth and demography in
 response to feral water buffalo in savannas of northern Australia: an
 experimental field study in Kakadu National Park. Australian journal of
 botany 54:283–296. Westgate, M. J. 2019. revtools: An
 R package to support article screening for evidence synthesis.
 Wigley, B. J., D. J. Augustine, C. Coetsee, J. Ratnam, and M.
 Sankaran. 2020. Grasses continue to trump trees at soil carbon
 sequestration following herbivore exclusion in a semiarid African savanna.
 Ecology 101:e03008. Wilson, D. J., W. A. Ruscoe, L.
 E. Burrows, L. M. McElrea, and D. Choquenot. 2006. An experimental study
 of the impacts of understorey forest vegetation and herbivory by red deer
 and rodents on seedling establishment and species composition in Waitutu
 Forest, New Zealand. New Zealand journal of ecology:191–207.
 Wood, G. W., M. T. Mengak, and M. Murphy. 1987. Ecological
 importance of feral ungulates at Shackleford Banks, North Carolina. The
 American midland naturalist:236–244. Young, T. P., T.
 M. Palmer, and M. E. Gadd. 2005. Competition and compensation among
 cattle, zebras, and elephants in a semi-arid savanna in Laikipia, Kenya.
 Biological conservation 122:351–359. Zalba, S. M.,
 and N. C. Cozzani. 2004. The impact of feral horses on grassland bird
 communities in Argentina. Pages 35–44. Cambridge University
 Press. Zamin, T. J., and P. Grogan. 2013. Caribou
 exclusion during a population low increases deciduous and evergreen shrub
 species biomass and nitrogen pools in low A rctic tundra. The Journal of
 ecology 101:671–683. Zanne, A. E., D. C. Tank, W. K.
 Cornwell, J. M. Eastman, S. A. Smith, R. G. FitzJohn, D. J. McGlinn, B. C.
 O’Meara, A. T. Moles, P. B. Reich, D. L. Royer, D. E. Soltis, P. F.
 Stevens, M. Westoby, I. J. Wright, L. Aarssen, R. I. Bertin, A. Calaminus,
 R. Govaerts, F. Hemmings, M. R. Leishman, J. Oleksyn, P. S. Soltis, N. G.
 Swenson, L. Warman, and J. M. Beaulieu. 2015. Three keys to the radiation
 of angiosperms into freezing environments. Nature 521:380.
 Zhao, M., F. A. Heinsch, R. R. Nemani, and S. W. Running. 2005.
 Improvements of the MODIS terrestrial gross and net primary production
 global data set. Remote Sensing of the Environment 95:164–176.
description: R version 4.2.1 as well as a variety of packages, including
 data.table, metafor, multcomp, ggplot2, broom, tidyr, and dplyr.
description: # Data for Functional traits - not nativeness - shape the effects of large
 mammalian herbivores on plant communities --- This dataset consists of
 meta-analytic data on plant diversity and abundance responses to mammalian
 megafauna (&gt;45 kg). This data can be used (as in manuscript) to assess
 whether there are differences between introduced and native megafauna in
 their effects on plants and to assess how megafauna functional traits,
 environmental variables, and so on, influence megafauna effects on plants.
 ## Description of the Data and file structure This repository includes the
 following data files. Note that NA and blank cells are considered
 equivalent in all csv files. 'Plant_Abundance.csv' = Each row is
 an observed effect of megafauna on plant abundance.
 'Plant_Diversity.csv' = Each row is an observed effect of
 megafauna on plant diversity. This dataset and
 'Plant_Abundance.csv' have the same columns but are separated
 for convenience. 'Model_Comparison_Guide.csv' = This dataset has
 a row for every model comparison used in manuscript. These comparisons
 were multi-tiered, comparing a null model to a base model (containing a
 factor of interest) and then this base model to a model with that factor
 of interest + nativeness. The filepath to each of these model objects is
 provided in the 'dir' columns ('dir_null',
 'dir_base', and 'dir_nativeness') and the formulas for
 each model are provided in the 'formula' columns.
 'models.zip' = This archive contains each rma.mv model object as
 an .Rds file, which can be loaded with readRDS() in R after loading the
 'metafor' package. 'Plant_Metadata.csv' = This csv
 file contains a description of each column name in the
 'Plant_Abundance.csv' and 'Plant_Diversity.csv'
 datasets as well as the variable names in all model objects (in
 'models.zip'), which themselves are trained on the data in
 'Plant_Abundance.csv' and 'Plant_Diversity.csv'
 'Model_Comparison_Metadata.csv' = This csv file contains a
 description of each column name in the
 'Model_Comparison_Guide.csv' file. The repository includes the
 following R scripts: 0_Helper_Scripts.R contains functions used in the
 other scripts. 1_Run_Models.R contains code to run all models. Note that
 this script should be used cautiously as some models can take days to run
 on a personal computer. 2_Analyze_and_Plot.R contains code to create
 figures and interpret models. ## Sharing/access Information Please cite:
 Lundgren et al. Functional traits - not nativeness - shape the effects of
 large mammalian herbivores on plant communities. Science.