Data from: Release of tree species diversity follows loss of elephants from evergreen tropical forests
Data files
Apr 27, 2026 version files 473.66 KB
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1._Seedling_plots.csv
10.68 KB
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2._Gabon_sapling.csv
15.52 KB
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3._Malaysia_sapling.csv
53.99 KB
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4._Gabon_tree.csv
4.21 KB
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5._Gabon_preference.csv
1.59 KB
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6._Plots_Malaysia.csv
139.92 KB
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7._Diet_Malaysia.csv
31.28 KB
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8._Impact_Malaysia.csv
86.63 KB
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9._Availability_plots_Malaysia.csv
110.08 KB
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Readme_Terbough_et_al._2025.txt
2.84 KB
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README.md
16.90 KB
Abstract
Little is known about the impacts of megaherbivores in closed-canopy tropical evergreen forests. We report on a decade of research on megaherbivore impacts in such forests in Gabon and Malaysia, comparing sites with (+) and without (-) elephants and documenting major differences in forest structure, tree species composition, and tree species diversity. In both regions, we compared sites supporting natural densities of elephants with otherwise undisturbed sites from which elephants had been absent for several decades. Elephant (+) sites supported low densities of seedlings and saplings relative to elephant (-) sites. In Lope National Park, Gabon, 88% of saplings and small trees (<20 cm dbh) were of species avoided by elephants, implicating forest elephants as powerful filters in tree recruitment. In Malaysia, Asian elephants showed strong preferences for monocots over dicots, as we found through both indirect and direct means. Loss of elephants from both Asian and African forests releases diversity from top-down pressure, as preferred forage species increase in abundance, leading to increased density of small stems and increased tree species diversity. In contrast, loss of other major functional groups of animals, including top carnivores, seed predators and seed dispersers, often results in negative impacts on tree diversity.
The dataset is collected from direct observations of elephant feeding and indirect observations through plot measurements from Gabon, Malaysia and Peru. It combines data from authors of the following studies for our comparative analysis:
- Terborgh J, Davenport LC, Niangadouma R, Dimoto E, Mouandza JC, Schultz O, Jaen MR. 2016 The African rainforest: odd man out or megafaunal landscape? African and Amazonian forests compared. Ecography 39, 187–193. (doi:10.1111/ecog.01643)
- White LJT. 1994 Biomass of Rain Forest Mammals in the Lope Reserve, Gabon. J. Anim. Ecol. 63, 499. (doi:10.2307/5217)
- Terborgh J, Davenport LC, Ong L, Campos‐Arceiz A. 2018 Foraging impacts of Asian megafauna on tropical rain forest structure and biodiversity. Biotropica 50, 84–89. (doi:10.1111/btp.12488)
- Ong, L., Tan, W. H., Davenport, L. C., McConkey, K. R., Mat Amin, M. K. A. B., Campos-Arceiz, A., & Terborgh, J. W. (2023). Asian elephants as ecological filters in Sundaic forests. Frontiers in Forests and Global Change, 6, 1143633. (doi.org/10.3389/ffgc.2023.1143633)
Abbreviations
- dbh: diameter at breast height
- ht.: height
Legend for Categorial Variables
- Plant.Class / group: monocot Monocotyledonous plant, such as grasses, palms, herbs, or other monocots.
- Plant.Class / group: dicot Dicotyledonous plant, such as most trees and lianas.
- Lifeform: tree- Woody tree stem.
- Lifeform: liana- Woody climbing plant.
- Lifeform: grass- Grass life form.
- Lifeform: herb- Non-woody herbaceous plant.
- Lifeform: palm- Palms.
- Plant.Size / plant.structure: below 1m tall- Seedlings or stem below 1 m in height.
- Plant.Size / plant.structure: small saplings- Small Sapling size class ≥1 m tall and ≤1 cm dbh.
- Plant.Size / plant.structure: large saplings- Large Sapling size class ≥1 cm and <10 cm dbh.
- Habitat.Type: Early successional- Early successional habitat or secondary growth habitat.
- Habitat.Type: Closed forest- Closed and mature forest.
- Patches: foraged- Patch where elephant foraging occurred.
- Patches: unforaged- Patch where no elephant foraging occurred.
- Eaten.or.Intact: eaten- Plant that showed evidence of elephant feeding or handling.
- Eaten.or.Intact: intact- Plant that did not show feeding or handling damage.
- Intact, Eaten, Mainstem.broken, Broken.Uprooted.Debarked: 1 Condition is present / true.
- Intact, Eaten, Mainstem.broken, Broken.Uprooted.Debarked: 0 Condition is absent / false.
- Herbivory.handling / Plant.handling / Herbivory: uproot / uprooted- Plant was uprooted during feeding.
- Herbivory.handling / Plant.handling / Herbivory: stem breakage / stem.breakage- Stem was broken during feeding.
- Herbivory.handling / Plant.handling / Herbivory: main stem breakage- Main stem was broken during feeding.
- Herbivory.handling / Plant.handling / Herbivory: leaves stripping- Leaves were stripped from the plant during feeding.
- Liana: yes- Recorded stem was a liana.
- Liana: no- Recorded stem was not a liana.
- Plant.Ht.below.1m: yes Plant height was below 1 m.
- Plant.Ht.below.1m: no Plant height was not below 1 m.
- Preference: strongly preferred- Species classified as strongly preferred by elephants.
- Preference: somewhat preferred- Species classified as somewhat preferred by elephants.
- Preference: avoided- Species classified as avoided by elephants.
- Preference: NA- No elephant preference category assigned; neither preferred or avoided.
- Species, Plant.Name, Plant_Local.name: taxon/local name Plant identity label, either scientific name, local name, genus-level ID, morphospecies, or unidentified label.
NA: Missing, unavailable, not recorded, or not applicable.
Filenames
1._Seedling_plots.csv – This file contains seedling plot data collected from sampling strips across different countries, including counts of seedlings recorded in each plot.
Variables Descriptions
- Plot ID: Unique plot identifier. For example, Lope 3.1.
- Location: Study location or site name (Lope, Mondah, Belum or Krau)
- Country Country: where the plot was sampled (Gabon, Peru or Malaysia)
- Quadrant: Sampling quadrant or strip within the plot, including cardinal directions SW, NW, NE, and SE.
- Trees_dbh10cm_ht.below1m: Number of tree sapling stems in the sampled strip with diameter at breast height less than 10 cm and height below 1 m.
- Lianas_dbh10cm_ht.below1m: Number of liana stems in the sampled strip with diameter at breast height less than 10 cm and height below 1 m.
- Herbs_dbh10cm_ht.below1m: Number of herbaceous stems in the sampled strip with diameter at breast height less than 10 cm and height below 1 m.
- Very.small_ht.below10cm: Number of very small stems or individuals with height below 10 cm.
- Unknown_ht.below15cm: Number of stems or individuals in which the plant type is unidentifiable and were below 15 cm tall.
- Unknown_ht.above15cm: Number of stems or individuals in which the plant type is unidentifiable and were above 15 cm tall.
- Columns.FandG.combined: Combined count from columns F (Lianas) and G (Herbs).
- All_ht.below1m: Total number of stems or individuals below 1 m in height recorded in the sampled strip. This is a summed total across the relevant stem-count categories.
2._Gabon_sapling.csv – This file records the abundance of small (SS) and large saplings (LS) in Gabon sampling strips.
Variables Descriptions
- SN: Serial number or row identifier for each species record.
- Species: Plant species name, or morphospecies level where species identification was not possible. Examples include Acepsia sp., Acindocarpus, Afrostyrax sp., Unknown sp. 1
- Mondah_1_SS: Stem count / Count of stems of the species recorded in Mondah strip 1 in the Small Sapling size class.
- Mondah_1_LS: Stem count in Mondah strip 1 in the Large Sapling size class.
- Mondah_2_SS: Stem count in Mondah strip 2 in the Small Sapling size class.
- Mondah_2_LS: Stem count in Mondah strip 2 in the Large Sapling size class.
- Mondah_3_SS: Stem count in Mondah strip 3 in the Small Sapling size class.
- Mondah_3_LS: Stem count in Mondah strip 3 in the Large Sapling size class.
- Mondah_4_SS: Stem count in Mondah strip 4 in the Small Sapling size class.
- Mondah_4_LS: Stem count in Mondah strip 4 in the Large Sapling size class.
- Mondah_5_SS: Stem count in Mondah strip 5 in the Small Sapling size class.
- Mondah_5_LS: Stem count in Mondah strip 5 in the Large Sapling size class.
- Lope_Strip_1_SS: Stem count in Lopé strip 1 in the Small Sapling size class.
- Lope_Strip_1_LS: Stem count in Lopé strip 1 in the Large Sapling size class.
- Lope_Strip_2_SS: Stem count in Lopé strip 2 in the Small Sapling size class.
- Lope_Strip_2_LS: Stem count in Lopé strip 2 in the Large Sapling size class.
- Lope_Strip_3_SS: Stem count in Lopé strip 3 in the Small Sapling size class.
- Lope_Strip_3_LS: Stem count in Lopé strip 3 in the Large Sapling size class.
- Lope_Strip_4_SS: Stem count in Lopé strip 4 in the Small Sapling size class.
- Lope_Strip_4_LS: Stem count in Lopé strip 4 in the Large Sapling size class.
- Lope_Strip_5_SS: Stem count in Lopé strip 5 in the Small Sapling size class.
- Lope_Strip_5_LS: Stem count in Lopé strip 5 in the Large Sapling size class.
3._Malaysia_sapling.csv – This file includes sapling abundance data from Malaysia sampling strips, showing counts of both small saplings (SS) and large saplings (LS).
Variables Descriptions
- Species Local plant name recorded during field surveys. These names may represent species, genera, morphospecies, or locally recognized plant types.
- Belum_1_SS: Stem count / Count of stems recorded in Belum strip 1 in the Small Sapling size class.
- Belum_2_SS: Stem count in Belum strip 2 in the Small Sapling size class.
- Belum_3_SS: Stem count in Belum strip 3 in the Small Sapling size class.
- Belum_4_SS: Stem count in Belum strip 4 in the Small Sapling size class.
- Belum_5_SS: Stem count in Belum strip 5 in the Small Sapling size class.
- Belum_6_SS: Stem count in Belum strip 6 in the Small Sapling size class.
- Belum_7_SS: Stem count in Belum strip 7 in the Small Sapling size class.
- Belum_1_LS: Stem count in Belum strip 1 in the Large Sapling size class.
- Belum_2_LS: Stem count in Belum strip 2 in the Large Sapling size class.
- Belum_3_LS: Stem count in Belum strip 3 in the Large Sapling size class.
- Belum_4_LS: Stem count in Belum strip 4 in the Large Sapling size class.
- Belum_5_LS: Stem count in Belum strip 5 in the Large Sapling size class.
- Belum_6_LS: Stem count in Belum strip 6 in the Large Sapling size class.
- Belum_7_LS: Stem count in Belum strip 7 in the Large Sapling size class.
- Krau_1_SS: Stem count in Krau strip 1 in the Small Sapling size class.
- Krau_2_SS: Stem count in Krau strip 2 in the Small Sapling size class.
- Krau_3_SS: Stem count in Krau strip 3 in the Small Sapling size class.
- Krau_4_SS: Stem count in Krau strip 4 in the Small Sapling size class.
- Krau_5_SS: Stem count in Krau strip 5 in the Small Sapling size class.
- Krau_6_SS: Stem count in Krau strip 6 in the Small Sapling size class.
- Krau_7_SS: Stem count in Krau strip 7 in the Small Sapling size class.
- Krau_1_LS: Stem count in Krau strip 1 in the Large Sapling size class.
- Krau_2_LS: Stem count in Krau strip 2 in the Large Sapling size class.
- Krau_3_LS: Stem count in Krau strip 3 in the Large Sapling size class.
- Krau_4_LS: Stem count in Krau strip 4 in the Large Sapling size class.
- Krau_5_LS: Stem count in Krau strip 5 in the Large Sapling size class.
- Krau_6_LS: Stem count in Krau strip 6 in the Large Sapling size class.
- Krau_7_LS: Stem count in Krau strip 7 in the Large Sapling size class.
4._Gabon_tree.csv – This file provides data on the abundance of adult trees in Gabon sampling strips, reported as counts of trees.
Variables Descriptions:
- Species: Scientific name of the tree. Some are identified to species level, while others only to genus, or morphospecies (uncertain category).
- Location: Sampling location name in Gabon. Angak_Lope refers to the Angak tree plot in Lopé National Park, and Mondah refers to the Mondah forest.
- Country: Country where the plot was sampled. For this dataset, all records are from Gabon.
- Tree: Count of tree stems of the species recorded at that location, sampled in standard 1-ha plots, stems ≥10 cm dbh.
5._Gabon_preference.csv – This file contains plant species abundance and elephant feeding preference in Gabon sampling strips, recorded as counts of plant species and if it is preferred.
Variables Descriptions:
- Species: Scientific name or identification label of the plant taxon. Some entries are identified to species level, while others are identified only to genus or morphospecies.
- Lope All Strips_SS: Count of stems of the species recorded across all Lopé sampling strips in the Small Sapling size class.\
- Lope All Strips_LS: Count of stems of the species recorded across all Lopé sampling strips in the Large Sapling size class.
- Lope Angak Trees dbh.below.20 cm: Count of tree stems of the species recorded in the Angak plot at Lopé in the small-tree size class. This corresponds to trees ≥10 cm and <20 cm dbh.
- Preference: Elephant foraging preference category for the species. Categories include strongly preferred, somewhat preferred, and avoided. NA means the species was not assigned a preference category in the available classification.
6._Plots_Malaysia.csv – This file includes data on plant abundance and trait measurements in Malaysia sampling strips, such as plant counts, plant height, DBH or circumference, and GPS coordinates of the sampling strips.
Variables Descriptions:
- SN: Serial number for each recorded stem.
- Site: Study site where the stem was recorded in Belum, Malaysia.
- Plot ID: Plot identifier within the site.
- Plant.Ht_m: Height of the recorded plant stem in meters. NA indicates that height was not recorded.
- Dbh_mm: Diameter at breast height of the recorded stem, measured in millimeters. NA indicates that DBH was not recorded.
- Circumference_mm: Stem circumference measured in millimeters. NA indicates that circumference was not recorded.
- Liana: Indicates whether the recorded stem was a liana. Values are yes or no.
- Plant_Local.name: Local plant name recorded during field surveys. These names represent morphospecies, or locally recognized plant types.
- Location GPS coordinates of the plot or sampling location where the stem was recorded.
7._Diet_Malaysia.csv – This file documents diet observations of elephants in Malaysia, including the number of mouthfuls of different plant species eaten by individual elephants.
Variables Descriptions:
- SiteID: Sitet identifier where the elephant feeding observation was recorded.
- Habitat.Type: Habitat type where the feeding observation occurred.
- Observation.ID: Identifier for the elephant observation session.
- Elephant: Name or identifier of the observed elephant.
- Age: Age of the observed elephant, likely in years.
- Plant.Class: Broad plant class consumed or handled by the elephant (monocot or dicot).
- Lifeform: Growth form or life form of the plant consumed or handled (herb, grass, liana, palm, and tree).
- Plant.Size: Size class of the plant consumed or handled.
- Herbivory.handling: Type of elephant handling or feeding damage recorded.
- No.of.Mouthful.consumed: Number of mouthfuls consumed by the elephant for that plant category and handling type during the observation.
8._Impact_Malaysia.csv – This file records the impact of elephants on plants in Malaysia, including the number of stems handled and damaged by different elephants.
Variables Description
- unique ID: Unique row identifier for each recorded plant-handling event.
- Observers: Names of observers who recorded the elephant foraging event.
- Elephant.Observation.ID: Identifier for the elephant observation session.
- Transect.ID: Transect identifier where the observation was made. For example, D1P1.
- Habitat.Type: Habitat type where the elephant foraging observation occurred.
- Observation: Observation number within the elephant observation session.
- Mouth.ID: Identifier for the individual mouthful, feeding item, or sequential plant-handling record within an observation.
- Plant.size: Size class of the handled plant.
- Plant.Class: Broad plant class of the handled plant (monocot or dicot).
- Lifeform: Growth form or life form of the handled plant.
- Plant.handling: Type of handling or damage caused by the elephant while feeding.
- Plant.Ht.below.1m: Indicator of whether the handled plant or recorded plant height was below 1 m (yes or no).
9._Availability_plots_Malaysia.csv – This file describes plant availability and elephant impact in foraged patches in Malaysia, including data on available plant species, plant sizes, herbivory, and types of impact.
Variables Description
- statsID: Unique row identifier for each recorded plant stem.
- Plot ID: Plot identifier for the sampled patch. For example, D1P1F1.
- Site.ID: Site or transect identifier. For example, D1P1.
- Patches: Patch category, indicating whether the patch was foraged or not.
- Patch.ID: Identifier for the sampled patch within the plot or site. For example, H01.
- Habitat.Type: Habitat type where the patch was sampled.
- Plant ID: Unique plant identifier within the sampled patch.
- Plant.Name: Plant name or field identification label. This may be a local name, genus/species name, or NA if unidentified.
- Lifeform: Growth form of the recorded plant (tree, liana, palm, grass, or herb).
- group: Broad plant group (dicot or monocot).
- plant.structure: Plant size or structural class.
- Eaten.or.Intact: Categorical status indicating whether the plant was eaten/handled or remained intact.
- Intact: Binary indicator for whether the plant was intact. 1 = intact; 0 = not intact.
- Eaten: Binary indicator for whether the plant was eaten or handled by elephants. 1 = eaten/handled; 0 = not eaten/handled.
- Mainstem.broken: Binary indicator for whether the main stem was broken. 1 = main stem broken; 0 = main stem not broken.
- Broken.Uprooted.Debarked: Binary indicator for whether the plant showed major herbivory damage, including being broken, uprooted, or debarked. 1 = damaged; 0 = not damaged.
- Herbivory: Type of herbivory or damage recorded.
- Diameter_cm: Stem diameter of the plant, measured in centimeters.
- Break.diameter_cm: Diameter of the stem at the break point, measured in centimeters. NA indicates no break diameter was recorded.
- Break.height_m: Height of the break point above the ground, measured in meters. NA indicates no break height was recorded.
- Plant.height_m: Total plant height, measured in meters. NA indicates plant height was not recorded.
Datasets were collected from both direct elephant observations and indirect measurements of stems and past breaks (scars) from forest plots in Gabon and Malaysia, as well as forest plots measurements in Peru.
