Human-induced downsizing of animal communities weakens trait matching between tropical plants and frugivores
Data files
Nov 14, 2025 version files 2.25 MB
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Analytical_workflow_Guerra_et_al._Ecol_Let.R
17.86 KB
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Data_Analytical_workflow_Guerra_et_al._Ecol_Let.RData
277.77 KB
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Dataset_S1_Network_Plant_Frugivore_Complete.xlsx
1.21 MB
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Dataset_S2_Fruit_and_Seed_Traits.xlsx
556.56 KB
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Dataset_S3_Frugivore_Traits.xlsx
104.54 KB
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Dataset_S4_Network_TraitMatch_Env_v2.xlsx
57.01 KB
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README.md
29.79 KB
Abstract
Defaunation of large-bodied animals threatens essential ecosystem functions, such as seed dispersal. However, the impact of this ‘downsizing’ of animal communities on plant-frugivore trait matching—the alignment of frugivory-related plant traits (e.g., fruit size) with frugivore traits (e.g., body mass)—remains poorly understood at macroecological scales. Here, we investigate how human disturbance and environmental conditions influence trait matching in plant-frugivore networks across the tropics. We compiled data on fruit size and several other frugivory-related plant traits, frugivore body mass, and diet information for 1,927 plant and 1,120 animal species and integrated these with 12,708 interaction records across 102 networks. The datasets provided include: Plant-frugivore interaction data (Dataset S1), the plant trait data (Dataset S2), the frugivore trait data (Dataset S3), and the network data (incl. environmental and human impact data; Dataset S4). Data sources are provided in the datasets, and each dataset contains a README sheet with additional information. Data is openly available for reuse.
Dataset DOI: 10.5061/dryad.g1jwstr3h
Description of the data and file structure
Four datasets are provided, including different columns with information.
Files and variables
File: Dataset_S1_Network_Plant_Frugivore_Complete.xlsx
Description: Plant-frugivore network data: Dataset S1.
We sourced the plant-frugivore interaction data from a comprehensive global dataset compiled by Fricke and Svenning (2020). This dataset consists of 410 local interaction networks, each constructed based on direct observations of animals interacting with fruits or seeds or from seeds retrieved from captured animals. These networks are bipartite, representing interactions between animal and plant species. While some networks recorded interaction frequencies, we used them in a binary format (presence or absence) due to inconsistencies in how frequencies were recorded across the dataset. This study focused on a subset of networks from tropical regions, defined geographically as the area between the Tropic of Cancer and the Tropic of Capricorn. This focus was chosen due to the key role of frugivorous vertebrates in seed dispersal within tropical ecosystems. The initial subset contained 251 spatially and temporally distinct tropical interaction networks (see Dataset S1). To account for repeated sampling at the same location over time, we aggregated unique plant-frugivore interactions from networks with the same coordinates into single composite networks, resulting in a final dataset of 181 geographically distinct networks (n = 145 non-composite networks; n = 36 composite networks formed by merging 2–11 original networks, mean ± SD: 2.94 ± 1.67). Prior to data analysis, the nomenclature of animal and plant species was updated to reflect taxonomic revisions made since the compilation of the original dataset. Animal species names were standardised following HBW and BirdLife International for birds, the International Union for Conservation of Nature (IUCN 2022) for mammals, and Oskyrko et al. (2024) for reptiles. Plant species names were standardised using the World Checklist of Vascular Plants (WCVP).
Variables
| Column Name | Description | Data Type | Units |
|---|---|---|---|
| animal.accepted.species | Accepted species name according to HBW and BirdLife International (2021) for birds, the International Union for Conservation of Nature (IUCN 2022) for mammals, and Oskyrko et al. (2024) for reptiles | Text | - |
| WCVP_plant_species_name | Accepted species name according to the World Checklist of Vascular Plants (WCVP) | Text | - |
| net.id | Unique identifier assigned to each ecological network (see Fricke and Svenning, 2020) | Text | - |
| study.id | Unique identifier assigned to each ecological network study (see Fricke and Svenning, 2020) | Text | - |
| latitude | Continuous | Decimal degrees (°) | |
| longitude | Continuous | Decimal degrees (°) | |
| locality | Name of the locality where the network was sampled | Text | - |
| country | Name of the country where the network was sampled | Text | - |
| year | Year in which the network was sampled | Continuous | - |
| Dispersal_unit_smaller_Axis | Measure of 'fruit size'. Measurement of the smallest axis of the dispersal unit, recorded as either fruit length or width for indehiscent fruits, or seed length or width for dehiscent fruits | Continuous | mm |
| BodyMass_g | Continuous | g | |
| Beak_volume | Continuous | mm3 | |
| Gape_Size | Continuous | mm | |
| Frugivory_level | Indicates the percentage of the diet that is made up of fruit | Continuous | - |
| Diet_Certainty | Specifies whether dietary information was inferred at the species, genus, or family level (see Wilman et al. 2014 for details) | Categorical | ABC, A, B, C, D1, D2 |
File: Dataset_S2_Fruit_and_Seed_Traits.xlsx
Description: Plant functional trait data: Dataset S2
We compiled data on frugivory-related plant traits, including fruit and seed dimensions (length and width), fruit type (e.g., berry, drupe, capsule; see Dataset S1), dehiscence (presence or absence), and seed fleshy appendages (presence or absence) for all 2,406 plant species that were present in the 181 ecological networks (see supplementary methods in Text S1 for details). Trait data were complete for 94.4% of the species (n = 2,272) and were primarily sourced from floras and monographs (79%), supplemented by scientific articles (19%) and photographs (2%) (see Dataset S2 for data and sources). Plant trait data were preferred under the accepted species name; however, when unavailable, they were retrieved from taxonomical homotypic synonyms following the WCVP. Methods for measuring fruit size vary considerably across studies. Some approaches consider fruit length or width, while others use a composite measure that sums fruit length and width. Both approaches present limitations: fruit length or width may not always correspond to the fruit’s longest or shortest axis, and combining length and width into a single metric may fail to capture the constraints frugivores face during consumption. To address these inconsistencies, we defined fruit size as the smallest reported axis of the dispersal unit (either fruit or seed), as this dimension is most likely to represent the key constraint for frugivores during ingestion. For indehiscent fruits, frugivores generally interact with the entire fruit, as the fleshy pericarp remains intact. In contrast, for dehiscent fruits, frugivores often interact with the seeds rather than the usually dry pericarp, especially when fleshy structures such as arils, sarcotestae, or elaiosomes are present. Accordingly, for the trait matching analyses, we selected the smallest axis of the dispersal unit, whether it was fruit length or width for indehiscent fruits or seed length or width for dehiscent fruits. Throughout the study, we refer to this measurement as ‘fruit size’.
Variables
Dataset S2: Plant trait data, fruit and seed traits
| Column Name | Description | Data Type | Units |
|---|---|---|---|
| Network_plant_species_name | Name under which the species appears in the original interaction networks | Text | - |
| WCVP_match_status | Indicates whether the name under which the species appears in the original interaction networks is accepted or a synonym | Text | - |
| WCVP_plant_species_name | Accepted species name according to the World Checklist of Vascular Plants (WCVP) | Text | - |
| Researcher | Researcher who collected and finalised trait data | Text | - |
| Mean_Fruit_length | Continuous | mm | |
| Min_Fruit_length | Continuous | mm | |
| Max_Fruit_length | Continuous | mm | |
| Mean_Fruit_width | Continuous | mm | |
| Min_Fruit_width | Continuous | mm | |
| Max_Fruit_width | Continuous | mm | |
| Mean_Seed_length | Continuous | mm | |
| Min_Seed_length | Continuous | mm | |
| Max_Seed_length | Continuous | mm | |
| Mean_Seed_width | Continuous | mm | |
| Min_Seed_width | Continuous | mm | |
| Max_Seed_width | Continuous | mm | |
| Fruit_Type | Describes the morphological type of the fruit (e.g., berry, drupe, capsule) | Text | - |
| Fruit_Dehiscence | Indicates whether the fruit splits open (dehiscent) or remains closed (indehiscent) at maturity | Binary | Dehiscent, Indehiscent |
| Seed_Fleshy_Appendage | Indicates whether the seed has a fleshy aril, sarcotesta, elaiosome or any other fleshy structure | Binary | Yes, No |
| Retrieved_from | Identifies the website or database from which the trait data was obtained | Text | - |
| URL | Provides the link to the website or database page containing the species’ trait data | Text | - |
| Source_Continuous | Cites the backbone references used to score continuous trait data | Text | - |
| Source_Quality | Indicates the type (quality) of the backbone reference for the continuous trait data | Categorical | Flora, Monograph, Scientific Article, Photograph, Report, Website |
| Source_Categorical | Cites the backbone references used to score fruit type, dehiscence, and seed fleshy appendage | Text | - |
| Notes | Contains any additional remarks or notable details about the trait data | Text | - |
| Based_on_GenusAverage | Indicates whether the fruit or seed trait data was inferred using genus-level averages | Binary | Yes, No |
File: Dataset_S3_Frugivore_Traits.xlsx
Description: Frugivore functional trait data: Dataset S3
We compiled data on frugivory-related animal traits, including body mass, degree of frugivory (percentage of the diet composed of fruit), beak dimensions, and gape size for all 1,226 animal species that were present in the 181 ecological networks. Trait data were complete for 98.9% of the species (n = 1,213) and were primarily sourced from established databases: AVONET, COMBINE, EltonTraits, ReptTraits (see Dataset S3 for data and sources). Animal trait data were preferred under the accepted species name; however, when unavailable, they were retrieved from taxonomic synonyms, following HBW and BirdLife International for birds and Soria et al. (2021) for mammals. In cases of discrepancies between accepted names and synonyms, the mean trait values were used. Dietary information was available for 1,102 species, with species-specific data for 93.7% of them; for the remaining 6.3%, diet was inferred at the genus or family level (see Wilman et al. 2014 for details). We selected body mass as the primary trait because it provides a comparable metric across birds, mammals and reptiles. While bird-specific traits such as beak volume and gape size are often used in trait matching studies, body mass was chosen for its broader applicability across taxa. This decision was supported by the strong correlations observed between body mass and beak volume (r = 0.81) and gape size (r = 0.85) in birds. The final dataset was predominantly composed of birds (75.8%), followed by mammals (23.3%) and reptiles (0.8%).
Variables
Dataset S3: Animal trait data, frugivore traits
| Column Name | Description | Data Type | Units |
|---|---|---|---|
| All_Network_Names | Name under which the species appears in the original interaction networks | Text | - |
| animal.class | Text | - | |
| animal.accepted.species | Accepted species name according to HBW and BirdLife International (2021) for birds, the International Union for Conservation of Nature (IUCN 2022) for mammals, and Oskyrko et al. (2024) for reptiles | Text | - |
| BodyMass_g | Continuous | g | |
| Beak_volume | Continuous | mm3 | |
| Gape_Size | Continuous | mm | |
| Frugivory_level | Indicates the percentage of the diet that is made up of fruit | Continuous | - |
| Granivory_level | Indicates the percentage of the diet that is made up of seed | Continuous | - |
| Diet_Certainty | Specifies whether dietary information was inferred at the species, genus, or family level (see Wilman et al. 2014 for details) | Categorical | ABC, A, B, C, D1, D2 |
| Body_Mass_Sources | Identifies the data sources used for body mass | Text | - |
| Beak_Volume_Sources | Identifies the data source used for beak volume | Text | - |
| Gape_Size_Sources | Identifies the data source used for gape size | Text | - |
| Diet_Sources | Identifies the data sources used for the percentage of the diet that is made up of fruit | Text | - |
| Notes | Contains any additional remarks or notable details about the trait data | Text | - |
File: Dataset_S4_Network_TraitMatch_Env_v2.xlsx
Description:
Human disturbance and environmental data: Dataset S4
Human disturbance and environmental data were assembled and extracted for each plant-frugivore network at spatial resolutions of 1 km and 5 km using ArcGIS Pro 3.3. Human disturbance data were derived from the Global Human Modification (gHM) dataset (Kennedy et al. 2020), which provides an integrated (i.e., cumulative) measure of human alteration of terrestrial landscapes at 1 km resolution. This continuous metric (ranging from 0 to 1) represents the proportion of land modified by human activities and integrates 13 indicators of disturbance, including human population density, built-up areas, cropland, livestock, major roads, minor roads, two-track roads, railroads, mining, oil wells, wind turbines, powerlines, and night-time lights, based on spatially explicit global datasets with a median reference year of 2016 (Kennedy et al. 2020). Environmental data were sourced from CHELSA V2.1, which provided annual temperature (°C), precipitation (mm), and NPP (g C/m²) at a 1 km resolution, averaged over the period 1981–2010. Although precipitation and NPP showed a strong correlation (r = 0.71), this remained below the commonly accepted multicollinearity threshold of r = 0.80. Consequently, both variables were retained, as their influence on frugivory may act through distinct ecological mechanisms. Additionally, AET (mm) was extracted as the annual average across the period 1982–2019 at a 1 km resolution. All network-level data is provided in Dataset S4.
Variables
Dataset S4: Environmental and human impact data per network
| Column Name | Description | Data Type | Units |
|---|---|---|---|
| net.id | Unique identifier assigned to each ecological network (see Fricke and Svenning, 2020) | Text | - |
| weighted_z | Trait matching between plant fruit size and frugivore body mass, calculated from all unique species interactions in plant-frugivore networks | Continuous | Weighted z-transformed Pearson’s r |
| latitude | Continuous | Decimal degrees (°) | |
| longitude | Continuous | Decimal degrees (°) | |
| year | Year in which the network was sampled | Continuous | - |
| sampling_method | The sampling method used to record interactions between plants and frugivores | Text | - |
| biogeographical_realm | The biogeographic realm where the network is located | Text | - |
| animal_richness | Total number of distinct animal species present in the network | Continuous | - |
| plant_richness | Total number of distinct plant species present in the network | Continuous | - |
| interaction_count | Total number of unique recorded interactions between plants and frugivores in the network | Continuous | - |
| avg_frugivory_per_site | Average degree of frugivory across all plant-frugivore interactions | Continuous | - |
| fruit_size_range | Range (max-min) of the log-transformed fruit sizes within the network | Continuous | - |
| body_mass_range | Range (max-min) of the log-transformed frugivore body masses within the network | Continuous | - |
| mean_annual_precipitation | Mean annual precipitation at 1 km resolution, averaged over 1981–2010 | Continuous | mm |
| mean_annual_temperature | Mean annual temperature at 1 km resolution, averaged over 1981–2010 | Continuous | °C |
| net_primary_productivity | Mean annual net primary productivity at 1 km resolution, averaged over 1981–2010 | Continuous | g C/m² |
| mean_annual_evapotranspiration | Mean annual actual evapotranspiration at 1 km resolution, averaged over 1982–2019 | Continuous | mm |
| ghm | Index of Global Human Modification (gHM) intensity (Kennedy et al. 2020) | Continuous | - |
Code/software
Analytical workflow to reproduce main results from Structural Equation Models and Figure 3 are provided in the following files, including the R script and the R data to be loaded into R.
- Analytical_workflow_Guerra_et_al._Ecol_Let.R
- Data_Analytical_workflow_Guerra_et_al._Ecol_Let.RData
Access information
Other publicly accessible locations of the data:
- In the supporting information of the article.
Data was derived from the following sources:
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Fricke, E.C. & Svenning, J.-C. (2020). Accelerating homogenization of the global plant–frugivore meta-network. Nature, 585, 74–78.
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Karger, D.N., Conrad, O., Böhner, J., Kawohl, T., Kreft, H., Soria-Auza, R.W., et al. (2017). Climatologies at high resolution for the earth’s land surface areas. Sci Data, 4, 170122.
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Kennedy, C.M., Oakleaf, J.R., Theobald, D.M., Baruch-Mordo, S. & Kiesecker, J. (2020). Global Human Modification of Terrestrial Systems.
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Oskyrko, O., Mi, C., Meiri, S. & Du, W. (2024). ReptTraits: a comprehensive dataset of ecological traits in reptiles. Sci Data, 11, 243.
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Soria, C.D., Pacifici, M., Di Marco, M., Stephen, S.M. & Rondinini, C. (2021). COMBINE: a coalesced mammal database of intrinsic and extrinsic traits. Ecology, 102, e03344.
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Wilman, H., Belmaker, J., Simpson, J., de la Rosa, C., Rivadeneira, M.M. & Jetz, W. (2014). EltonTraits 1.0: Species-level foraging attributes of the world’s birds and mammals. Ecology, 95, 2027–2027.
