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Fulltext The first record of a fly of the family Milichiidae (Diptera) interacting with an ant of the genus Polyrhachis Smith, 1857 (Hymenoptera: Formicidae)

Yusah KM, Fayle TM

Biodivers Data J, 2014.
PMID: 25425942 DOI: 10.3897/BDJ.2.e4168

Flies in the family Milichiidae are often myrmecophilic. We document the first record of a fly from this family interacting with an ant of the genus Polyrhachis. In lowland riparian rainforest in Sabah, Malaysia, we observed a female of the genus Milichia following an ant of the species of P.illaudata, and repeatedly attempting to make close contact. Our observation suggests that the dipteran may have been attempting to feed kleptoparasitically from the Polyrhachis worker, since members of this ant genus often feed on liquid carbohydrate-rich food resources. This is the first time an interaction has been observed between a fly of this family and an ant of this widespread old world tropical genus.
Ant-termite interactions: an important but under-explored ecological linkage

Tuma J, Eggleton P, Fayle TM

Biol Rev Camb Philos Soc, 2020 06;95(3):555-572.
PMID: 31876057 DOI: 10.1111/brv.12577

Animal interactions play an important role in understanding ecological processes. The nature and intensity of these interactions can shape the impacts of organisms on their environment. Because ants and termites, with their high biomass and range of ecological functions, have considerable effects on their environment, the interaction between them is important for ecosystem processes. Although the manner in which ants and termites interact is becoming increasingly well studied, there has been no synthesis to date of the available literature. Here we review and synthesise all existing literature on ant-termite interactions. We infer that ant predation on termites is the most important, most widespread, and most studied type of interaction. Predatory ant species can regulate termite populations and subsequently slow down the decomposition of wood, litter and soil organic matter. As a consequence they also affect plant growth and distribution, nutrient cycling and nutrient availability. Although some ant species are specialised termite predators, there is probably a high level of opportunistic predation by generalist ant species, and hence their impact on ecosystem processes that termites are known to provide varies at the species level. The most fruitful future research direction will be to evaluate the impact of ant-termite predation on broader ecosystem processes. To do this it will be necessary to quantify the efficacy both of particular ant species and of ant communities as a whole in regulating termite populations in different biomes. We envisage that this work will require a combination of methods, including DNA barcoding of ant gut contents along with field observations and exclusion experiments. Such a combined approach is necessary for assessing how this interaction influences entire ecosystems.
Riparian reserves within oil palm plantations conserve logged forest leaf litter ant communities and maintain associated scavenging rates

Gray CL, Lewis OT, Chung AY, Fayle TM

The Journal of applied ecology, 2015 Feb;52(1):31-40.
PMID: 25678717

The expansion of oil palm plantations at the expense of tropical forests is causing declines in many species and altering ecosystem functions. Maintaining forest-dependent species and processes in these landscapes may therefore limit the negative impacts of this economically important industry. Protecting riparian vegetation may be one such opportunity; forest buffer strips are commonly protected for hydrological reasons, but can also conserve functionally important taxa and the processes they support.We surveyed leaf litter ant communities within oil palm-dominated landscapes in Sabah, Malaysia, using protein baits. As the scavenging activity of ants influences important ecological characteristics such as nutrient cycling and soil structure, we quantified species-specific rates of bait removal to examine how this process may change across land uses and establish which changes in community structure underlie observed shifts in activity.Riparian reserves had similar ant species richness, community composition and scavenging rates to nearby continuous logged forest. Reserve width and vegetation structure did not affect ant species richness significantly. However, the number of foraging individuals decreased with increasing reserve width, and scavenging rate increased with vegetation complexity.Oil palm ant communities were characterized by significantly lower species richness than logged forest and riparian reserves and also by altered community composition and reduced scavenging rates.Reduced scavenging activity in oil palm was not explained by a reduction in ant species richness, nor by replacement of forest ant species by those with lower per species scavenging rates. There was also no significant effect of land use on the scavenging activity of the forest species that persisted in oil palm. Rather, changes in scavenging activity were best explained by a reduction in the mean rate of bait removal per individual ant across all species in the community.Synthesis and applications. Our results suggest that riparian reserves are comparable to areas of logged forest in terms of ant community composition and ant-mediated scavenging. Hence, in addition to protecting large continuous areas of primary and logged forest, maintaining riparian reserves is a successful strategy for conserving leaf litter ants and their scavenging activities in tropical agricultural landscapes.
Fulltext Ant mosaics in Bornean primary rain forest high canopy depend on spatial scale, time of day, and sampling method

Yusah KM, Foster WA, Reynolds G, Fayle TM

PeerJ, 2018;6:e4231.
PMID: 29423344 DOI: 10.7717/peerj.4231

Background: Competitive interactions in biological communities can be thought of as giving rise to "assembly rules" that dictate the species that are able to co-exist. Ant communities in tropical canopies often display a particular pattern, an "ant mosaic", in which competition between dominant ant species results in a patchwork of mutually exclusive territories. Although ant mosaics have been well-documented in plantation landscapes, their presence in pristine tropical forests remained contentious until recently. Here we assess presence of ant mosaics in a hitherto under-investigated forest stratum, the emergent trees of the high canopy in primary tropical rain forest, and explore how the strength of any ant mosaics is affected by spatial scale, time of day, and sampling method.
Methods: To test whether these factors might impact the detection of ant mosaics in pristine habitats, we sampled ant communities from emergent trees, which rise above the highest canopy layers in lowland dipterocarp rain forests in North Borneo (38.8-60.2 m), using both baiting and insecticide fogging. Critically, we restricted sampling to only the canopy of each focal tree. For baiting, we carried out sampling during both the day and the night. We used null models of species co-occurrence to assess patterns of segregation at within-tree and between-tree scales.
Results: The numerically dominant ant species on the emergent trees sampled formed a diverse community, with differences in the identity of dominant species between times of day and sampling methods. Between trees, we found patterns of ant species segregation consistent with the existence of ant mosaics using both methods. Within trees, fogged ants were segregated, while baited ants were segregated only at night.
Discussion: We conclude that ant mosaics are present within the emergent trees of the high canopy of tropical rain forest in Malaysian Borneo, and that sampling technique, spatial scale, and time of day interact to determine observed patterns of segregation. Restricting sampling to only emergent trees reveals segregatory patterns not observed in ground-based studies, confirming previous observations of stronger segregation with increasing height in the canopy.
Fulltext Tropical land-use change alters trait-based community assembly rules for dung beetles and birds

Edwards FA, Edwards DP, Hamer KC, Fayle TM

Oecologia, 2021 Mar;195(3):705-717.
PMID: 33559003 DOI: 10.1007/s00442-020-04829-z

Tropical rainforest disturbance and conversion are critical drivers of biodiversity loss. A key knowledge gap is understanding the impacts of habitat modification on mechanisms of community assembly, which are predicted to respond differently between taxa and across spatial scales. We use a null model approach to detect trait assembly of species at local- and landscape-scales, and then subdivide communities with different habitat associations and foraging guilds to investigate whether the detection of assembly mechanisms varies between groups. We focus on two indicator taxa, dung beetles and birds, across a disturbance gradient of primary rainforest, selectively logged rainforest, and oil palm plantations in Borneo, Southeast Asia. Random community assembly was predominant for dung beetles across habitats, whereas trait convergence, indicative of environmental filtering, occurred across the disturbance gradient for birds. Assembly patterns at the two spatial scales were similar. Subdividing for habitat association and foraging guild revealed patterns hidden when focusing on the overall community. Dung beetle forest specialists and habitat generalists showed opposing assembly mechanisms in primary forest, community assembly of habitat generalists for both taxa differed with disturbance intensity, and insectivorous birds strongly influenced overall community assembly relative to other guilds. Our study reveals the sensitivity of community assembly mechanisms to anthropogenic disturbance via a shift in the relative contribution of stochastic and deterministic processes. This highlights the need for greater understanding of how habitat modification alters species interactions and the importance of incorporating species' traits within assessments.
Fulltext The conservation value of South East Asia's highly degraded forests: evidence from leaf-litter ants

Woodcock P, Edwards DP, Fayle TM, Newton RJ, Khen CV, Bottrell SH, et al.

Philos Trans R Soc Lond B Biol Sci, 2011 Nov 27;366(1582):3256-64.
PMID: 22006966 DOI: 10.1098/rstb.2011.0031

South East Asia is widely regarded as a centre of threatened biodiversity owing to extensive logging and forest conversion to agriculture. In particular, forests degraded by repeated rounds of intensive logging are viewed as having little conservation value and are afforded meagre protection from conversion to oil palm. Here, we determine the biological value of such heavily degraded forests by comparing leaf-litter ant communities in unlogged (natural) and twice-logged forests in Sabah, Borneo. We accounted for impacts of logging on habitat heterogeneity by comparing species richness and composition at four nested spatial scales, and examining how species richness was partitioned across the landscape in each habitat. We found that twice-logged forest had fewer species occurrences, lower species richness at small spatial scales and altered species composition compared with natural forests. However, over 80 per cent of species found in unlogged forest were detected within twice-logged forest. Moreover, greater species turnover among sites in twice-logged forest resulted in identical species richness between habitats at the largest spatial scale. While two intensive logging cycles have negative impacts on ant communities, these degraded forests clearly provide important habitat for numerous species and preventing their conversion to oil palm and other crops should be a conservation priority.
Fulltext Whole-ecosystem experimental manipulations of tropical forests

Fayle TM, Turner EC, Basset Y, Ewers RM, Reynolds G, Novotny V

Trends Ecol Evol, 2015 Jun;30(6):334-46.
PMID: 25896491 DOI: 10.1016/j.tree.2015.03.010

Tropical forests are highly diverse systems involving extraordinary numbers of interactions between species, with each species responding in a different way to the abiotic environment. Understanding how these systems function and predicting how they respond to anthropogenic global change is extremely challenging. We argue for the necessity of 'whole-ecosystem' experimental manipulations, in which the entire ecosystem is targeted, either to reveal the functioning of the system in its natural state or to understand responses to anthropogenic impacts. We survey the current range of whole-ecosystem manipulations, which include those targeting weather and climate, nutrients, biotic interactions, human impacts, and habitat restoration. Finally we describe the unique challenges and opportunities presented by such projects and suggest directions for future experiments.
Fulltext Detection of mitochondrial COII DNA sequences in ant guts as a method for assessing termite predation by ants

Fayle TM, Scholtz O, Dumbrell AJ, Russell S, Segar ST, Eggleton P

PLoS One, 2015;10(4):e0122533.
PMID: 25853549 DOI: 10.1371/journal.pone.0122533

Termites and ants contribute more to animal biomass in tropical rain forests than any other single group and perform vital ecosystem functions. Although ants prey on termites, at the community level the linkage between these groups is poorly understood. Thus, assessing the distribution and specificity of ant termitophagy is of considerable interest. We describe an approach for quantifying ant-termite food webs by sequencing termite DNA (cytochrome c oxidase subunit II, COII) from ant guts and apply this to a soil-dwelling ant community from tropical rain forest in Gabon. We extracted DNA from 215 ants from 15 species. Of these, 17.2 % of individuals had termite DNA in their guts, with BLAST analysis confirming the identity of 34.1 % of these termites to family level or better. Although ant species varied in detection of termite DNA, ranging from 63 % (5/7; Camponotus sp. 1) to 0 % (0/7; Ponera sp. 1), there was no evidence (with small sample sizes) for heterogeneity in termite consumption across ant taxa, and no evidence for species-specific ant-termite predation. In all three ant species with identifiable termite DNA in multiple individuals, multiple termite species were represented. Furthermore, the two termite species that were detected on multiple occasions in ant guts were in both cases found in multiple ant species, suggesting that ant-termite food webs are not strongly compartmentalised. However, two ant species were found to consume only Anoplotermes-group termites, indicating possible predatory specialisation at a higher taxonomic level. Using a laboratory feeding test, we were able to detect termite COII sequences in ant guts up to 2 h after feeding, indicating that our method only detects recent feeding events. Our data provide tentative support for the hypothesis that unspecialised termite predation by ants is widespread and highlight the use of molecular approaches for future studies of ant-termite food webs.
Fulltext Biodiversity hanging by a thread: the importance of fungal litter-trapping systems in tropical rainforests

Snaddon JL, Turner EC, Fayle TM, Khen CV, Eggleton P, Foster WA

Biol Lett, 2012 Jun 23;8(3):397-400.
PMID: 22188674 DOI: 10.1098/rsbl.2011.1115

The exceptionally high species richness of arthropods in tropical rainforests hinges on the complexity of the forest itself: that is, on features such as the high plant diversity, the layered nature of the canopy and the abundance and the diversity of epiphytes and litter. We here report on one important, but almost completely neglected, piece of this complex jigsaw-the intricate network of rhizomorph-forming fungi that ramify through the vegetation of the lower canopy and intercept falling leaf litter. We show that this litter-trapping network is abundant and intercepts substantial amounts of litter (257.3 kg ha(-1)): this exceeds the amount of material recorded in any other rainforest litter-trapping system. Experimental removal of this fungal network resulted in a dramatic reduction in both the abundance (decreased by 70.2 ± 4.1%) and morphospecies richness (decreased by 57.4 ± 5.1%) of arthropods. Since the lower canopy levels can contain the highest densities of arthropods, the proportion of the rainforest fauna dependent on the fungal networks is likely to be substantial. Fungal litter-trapping systems are therefore a crucial component of habitat complexity, providing a vital resource that contributes significantly to rainforest biodiversity.
Fulltext Experimentally testing and assessing the predictive power of species assembly rules for tropical canopy ants

Fayle TM, Eggleton P, Manica A, Yusah KM, Foster WA

Ecol Lett, 2015 Mar;18(3):254-62.
PMID: 25622647 DOI: 10.1111/ele.12403

Understanding how species assemble into communities is a key goal in ecology. However, assembly rules are rarely tested experimentally, and their ability to shape real communities is poorly known. We surveyed a diverse community of epiphyte-dwelling ants and found that similar-sized species co-occurred less often than expected. Laboratory experiments demonstrated that invasion was discouraged by the presence of similarly sized resident species. The size difference for which invasion was less likely was the same as that for which wild species exhibited reduced co-occurrence. Finally we explored whether our experimentally derived assembly rules could simulate realistic communities. Communities simulated using size-based species assembly exhibited diversities closer to wild communities than those simulated using size-independent assembly, with results being sensitive to the combination of rules employed. Hence, species segregation in the wild can be driven by competitive species assembly, and this process is sufficient to generate observed species abundance distributions for tropical epiphyte-dwelling ants.
The first record of the parasitic myrmecophilous caterpillar Liphyrabrassolis (Lepidoptera, Lycaenidae) inside Asian weaver ant (Oecophyllasmaragdina) nests in oil palm plantations

Dwi Advento A, Yusah KM, Salim H, Naim M, Caliman JP, Fayle TM

Biodivers Data J, 2022;10:e83842.
PMID: 36761604 DOI: 10.3897/BDJ.10.e83842

Asian weaver ants (Oecophyllasmaragdina) are an important biocontrol agent in agricultural habitats. We conducted surveys in oil palm plantations in Riau, Indonesia for an obligate myrmecophilous butterfly larvae, Liphyrabrassolis (Lepidoptera, Lycaenidae), that is known to consume weaver ant larvae in other habitat types. We found L.brassolis larvae in five of the twenty nests surveyed, with larval presence not being related to weaver ant nest size. We also observed L.brassolis larvae in a weaver ant mass rearing facility. This is the first report of L.brassolis from oil palm plantations and may have implications for the use of weaver ants as biological control agents.
On the Perils of Ignoring Evolution in Networks

Segar ST, Fayle TM, Srivastava DS, Lewinsohn TM, Lewis OT, Novotny V, et al.

Trends Ecol Evol, 2020 Oct;35(10):865-866.
PMID: 32854959 DOI: 10.1016/j.tree.2020.07.016
The Role of Evolution in Shaping Ecological Networks

Segar ST, Fayle TM, Srivastava DS, Lewinsohn TM, Lewis OT, Novotny V, et al.

Trends Ecol Evol, 2020 05;35(5):454-466.
PMID: 32294426 DOI: 10.1016/j.tree.2020.01.004

The structure of ecological networks reflects the evolutionary history of their biotic components, and their dynamics are strongly driven by ecoevolutionary processes. Here, we present an appraisal of recent relevant research, in which the pervasive role of evolution within ecological networks is manifest. Although evolutionary processes are most evident at macroevolutionary scales, they are also important drivers of local network structure and dynamics. We propose components of a blueprint for further research, emphasising process-based models, experimental evolution, and phenotypic variation, across a range of distinct spatial and temporal scales. Evolutionary dimensions are required to advance our understanding of foundational properties of community assembly and to enhance our capability of predicting how networks will respond to impending changes.
Fulltext Establishing the evidence base for maintaining biodiversity and ecosystem function in the oil palm landscapes of South East Asia

Foster WA, Snaddon JL, Turner EC, Fayle TM, Cockerill TD, Ellwood MD, et al.

Philos Trans R Soc Lond B Biol Sci, 2011 Nov 27;366(1582):3277-91.
PMID: 22006968 DOI: 10.1098/rstb.2011.0041

The conversion of natural forest to oil palm plantation is a major current threat to the conservation of biodiversity in South East Asia. Most animal taxa decrease in both species richness and abundance on conversion of forest to oil palm, and there is usually a severe loss of forest species. The extent of loss varies significantly across both different taxa and different microhabitats within the oil palm habitat. The principal driver of this loss in diversity is probably the biological and physical simplification of the habitat, but there is little direct evidence for this. The conservation of forest species requires the preservation of large reserves of intact forest, but we must not lose sight of the importance of conserving biodiversity and ecosystem processes within the oil palm habitat itself. We urgently need to carry out research that will establish whether maintaining diversity supports economically and ecologically important processes. There is some evidence that both landscape and local complexity can have positive impacts on biodiversity in the oil palm habitat. By intelligent manipulation of habitat complexity, it could be possible to enhance not only the number of species that can live in oil palm plantations but also their contribution to the healthy functioning of this exceptionally important and widespread landscape.
Fulltext Forests and Their Canopies: Achievements and Horizons in Canopy Science

Nakamura A, Kitching RL, Cao M, Creedy TJ, Fayle TM, Freiberg M, et al.

Trends Ecol Evol, 2017 06;32(6):438-451.
PMID: 28359572 DOI: 10.1016/j.tree.2017.02.020

Forest canopies are dynamic interfaces between organisms and atmosphere, providing buffered microclimates and complex microhabitats. Canopies form vertically stratified ecosystems interconnected with other strata. Some forest biodiversity patterns and food webs have been documented and measurements of ecophysiology and biogeochemical cycling have allowed analyses of large-scale transfer of CO2, water, and trace gases between forests and the atmosphere. However, many knowledge gaps remain. With global research networks and databases, and new technologies and infrastructure, we envisage rapid advances in our understanding of the mechanisms that drive the spatial and temporal dynamics of forests and their canopies. Such understanding is vital for the successful management and conservation of global forests and the ecosystem services they provide to the world.
Fulltext Midpoint attractors and species richness: Modelling the interaction between environmental drivers and geometric constraints

Colwell RK, Gotelli NJ, Ashton LA, Beck J, Brehm G, Fayle TM, et al.

Ecol Lett, 2016 09;19(9):1009-22.
PMID: 27358193 DOI: 10.1111/ele.12640

We introduce a novel framework for conceptualising, quantifying and unifying discordant patterns of species richness along geographical gradients. While not itself explicitly mechanistic, this approach offers a path towards understanding mechanisms. In this study, we focused on the diverse patterns of species richness on mountainsides. We conjectured that elevational range midpoints of species may be drawn towards a single midpoint attractor - a unimodal gradient of environmental favourability. The midpoint attractor interacts with geometric constraints imposed by sea level and the mountaintop to produce taxon-specific patterns of species richness. We developed a Bayesian simulation model to estimate the location and strength of the midpoint attractor from species occurrence data sampled along mountainsides. We also constructed midpoint predictor models to test whether environmental variables could directly account for the observed patterns of species range midpoints. We challenged these models with 16 elevational data sets, comprising 4500 species of insects, vertebrates and plants. The midpoint predictor models generally failed to predict the pattern of species midpoints. In contrast, the midpoint attractor model closely reproduced empirical spatial patterns of species richness and range midpoints. Gradients of environmental favourability, subject to geometric constraints, may parsimoniously account for elevational and other patterns of species richness.
Fulltext Dispatch from the field: ecology of ground-web-building spiders with description of a new species (Araneae, Symphytognathidae)

Miller JA, Schilthuizen M, Burmester JL, van der Graaf L, Merckx V, Jocqué M, et al.

Biodivers Data J, 2014.
PMID: 24891829 DOI: 10.3897/BDJ.2.e1076

Crassignathadanaugirangensis sp. n. (Araneae: Symphytognathidae) was discovered during a tropical ecology field course held at the Danau Girang Field Centre in Sabah, Malaysia. A taxonomic description and accompanying ecological study were completed as course activities. To assess the ecology of this species, which belongs to the ground-web-building spider community, three habitat types were surveyed: riparian forest, recently inundated riverine forest, and oil palm plantation. Crassignathadanaugirangensis sp. n. is the most abundant ground-web-building spider species in riparian forest; it is rare or absent from the recently inundated forest and was not found in a nearby oil palm plantation. The availability of this taxonomic description may help facilitate the accumulation of data about this species and the role of inundated riverine forest in shaping invertebrate communities.
Fulltext Logging cuts the functional importance of invertebrates in tropical rainforest

Ewers RM, Boyle MJ, Gleave RA, Plowman NS, Benedick S, Bernard H, et al.

Nat Commun, 2015 Apr 13;6:6836.
PMID: 25865801 DOI: 10.1038/ncomms7836

Invertebrates are dominant species in primary tropical rainforests, where their abundance and diversity contributes to the functioning and resilience of these globally important ecosystems. However, more than one-third of tropical forests have been logged, with dramatic impacts on rainforest biodiversity that may disrupt key ecosystem processes. We find that the contribution of invertebrates to three ecosystem processes operating at three trophic levels (litter decomposition, seed predation and removal, and invertebrate predation) is reduced by up to one-half following logging. These changes are associated with decreased abundance of key functional groups of termites, ants, beetles and earthworms, and an increase in the abundance of small mammals, amphibians and insectivorous birds in logged relative to primary forest. Our results suggest that ecosystem processes themselves have considerable resilience to logging, but the consistent decline of invertebrate functional importance is indicative of a human-induced shift in how these ecological processes operate in tropical rainforests.
Fulltext Late-spring frost risk between 1959 and 2017 decreased in North America but increased in Europe and Asia

Zohner CM, Mo L, Renner SS, Svenning JC, Vitasse Y, Benito BM, et al.

Proc Natl Acad Sci U S A, 2020 06 02;117(22):12192-12200.
PMID: 32393624 DOI: 10.1073/pnas.1920816117

Late-spring frosts (LSFs) affect the performance of plants and animals across the world's temperate and boreal zones, but despite their ecological and economic impact on agriculture and forestry, the geographic distribution and evolutionary impact of these frost events are poorly understood. Here, we analyze LSFs between 1959 and 2017 and the resistance strategies of Northern Hemisphere woody species to infer trees' adaptations for minimizing frost damage to their leaves and to forecast forest vulnerability under the ongoing changes in frost frequencies. Trait values on leaf-out and leaf-freezing resistance come from up to 1,500 temperate and boreal woody species cultivated in common gardens. We find that areas in which LSFs are common, such as eastern North America, harbor tree species with cautious (late-leafing) leaf-out strategies. Areas in which LSFs used to be unlikely, such as broad-leaved forests and shrublands in Europe and Asia, instead harbor opportunistic tree species (quickly reacting to warming air temperatures). LSFs in the latter regions are currently increasing, and given species' innate resistance strategies, we estimate that ∼35% of the European and ∼26% of the Asian temperate forest area, but only ∼10% of the North American, will experience increasing late-frost damage in the future. Our findings reveal region-specific changes in the spring-frost risk that can inform decision-making in land management, forestry, agriculture, and insurance policy.
Fulltext Native diversity buffers against severity of non-native tree invasions

Delavaux CS, Crowther TW, Zohner CM, Robmann NM, Lauber T, van den Hoogen J, et al.

Nature, 2023 Sep;621(7980):773-781.
PMID: 37612513 DOI: 10.1038/s41586-023-06440-7

Determining the drivers of non-native plant invasions is critical for managing native ecosystems and limiting the spread of invasive species1,2. Tree invasions in particular have been relatively overlooked, even though they have the potential to transform ecosystems and economies3,4. Here, leveraging global tree databases5-7, we explore how the phylogenetic and functional diversity of native tree communities, human pressure and the environment influence the establishment of non-native tree species and the subsequent invasion severity. We find that anthropogenic factors are key to predicting whether a location is invaded, but that invasion severity is underpinned by native diversity, with higher diversity predicting lower invasion severity. Temperature and precipitation emerge as strong predictors of invasion strategy, with non-native species invading successfully when they are similar to the native community in cold or dry extremes. Yet, despite the influence of these ecological forces in determining invasion strategy, we find evidence that these patterns can be obscured by human activity, with lower ecological signal in areas with higher proximity to shipping ports. Our global perspective of non-native tree invasion highlights that human drivers influence non-native tree presence, and that native phylogenetic and functional diversity have a critical role in the establishment and spread of subsequent invasions.

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