Ecological traits of the world’s primates

Ecosystems largely depend, for both their functioning and their ecological integrity, on the ecological traits of the species that inhabit them. Non-human primates have a wide geographic distribution and play vital roles in ecosystem structure, function, and resilience. However, there is no comprehensive and updated compilation of information on ecological traits of all the world’s primate species to accurately assess such roles at a global scale. Here we present a database on some important ecological traits of the world’s primates (504 species), including home range size, locomotion type, diel activity, trophic guild, body mass, habitat type, current conservation status, population trend, and geographic realm. We compiled this information through a careful review of 1,216 studies published between 1941 and 2018, resulting in a comprehensive, easily accessible and user-friendly database. This database has broad applicability in primatological studies, and can potentially be used to address many research questions at all spatial scales, from local to global.

www.nature.com/scientificdata www.nature.com/scientificdata/ including function and resilience, and (6) the ecological roles of primates at different spatial scales. For example, we have used this database to test, at a global scale, which ecological traits of primates correlate more strongly with the use of the anthropogenic matrix in human-modified landscapes 10 .

Methods
Ecological traits included in the database are: home range size, locomotion type, diel activity, trophic guild, body mass, habitat type, current conservation status, population trend, and geographic realm. We selected these variables because of their well-known ecological significance 15 . For instance, body mass is correlated with many life-history traits that can affect the structure and dynamics of ecological networks 16,17 and is recognized as a variable that can have profound impact across multiple scales of organization, from the individual to the ecosystem level 16,[18][19][20] . We updated the primate taxonomic nomenclature following Estrada et al. 4 , which is mostly based on the International Union for Conservation of Nature (IUCN). Although we also indicate the most up-to-date primate taxonomic nomenclature published in the Integrated Taxonomic Information System (http://www.itis.gov/) of the Order Primates, we have probably not noticed the last taxonomic changes for some species. Thus, we kindly request users to contact the corresponding author if they find any error in the database to maintain it as updated as possible.
We collected data from 1,216 studies published between 1941 and 2018, including scientific articles, books, reports, dissertations, and web pages. The literature search included publications in English, Spanish, French, German and Portuguese. Geographic realm was extracted from Mittermeier et al. 3 Current conservation status, population trend and habitat type of each species were obtained from the IUCN database, using the 'letsR' 21 package for R, version 3.0.1. 22 . The IUCN defines habitat type as "the major habitats in which each taxon occurs" https://www.iucnredlist.org/resources/habitat-classification-scheme. When there was no information on these variables for a given primate species in the IUCN database, we used information available in other sources by actively searching in the World Wide Web (public domain and scientific publications). When different sources (2) locomotion types are terrestrial, both locomotion types, and arboreal; (3) diel activity includes diurnal, nocturnal and cathemeral; (4) trophic guild includes folivore, folivore-frugivore, frugivore, insectivore, omnivore, and gummivore (the latter not depicted); (5) body mass gradient from small to large; (6) habitat type includes seven categories (see text) but only two are depicted as examples (forest and savannah); (7) IUCN conservation status includes seven categories, with five depicted here (CR critically endangered, EN endangered, VU vulnerable, NT near threatened and LC least concern); (8) population trend is represented by three graphs indicating increasing, stable and decreasing populations; and (9) geographic realm is represented by a global map. Images used with permission from Microsoft.
www.nature.com/scientificdata www.nature.com/scientificdata/ yielded different information for a given species and trait, we recorded this information in different rows. This procedure allowed us to have a more comprehensive, accurate and objective database.
Information on home range sizes is given in hectares. When information and primary sources are available for the home range data and a study reports more than one home range size, we average all the values. We also include the minimum and maximum values, the number of groups and mean group size, the method used (e.g., minimum convex polygon), alternative home range and alternative method (when available), and the study duration. Locomotion type refers to the main way in which an animal moves in its environment. Arboreal locomotion type includes primate species that are strictly arboreal, which in undisturbed forest very rarely go to the ground; terrestrial type includes primate species that are mainly terrestrial, i.e., carrying out most of their daily activity on the ground; the locomotion category 'both' includes primate species which are commonly active on both substrates, ground as well as trees. Diel activity is categorized as diurnal (i.e., main behavioral activities occurring at daytime), nocturnal (i.e., main behavioral activities occurring at nighttime) and cathemeral (i.e., behavior occurring both at day and nighttime). In terms of trophic guild we consider six general groups: frugivore (>60% of fruits/seeds in diet), folivore (>60% leaves in diet), folivore-frugivore (diet comprised of both fruits/seeds and leaves in similar proportions), omnivore (diet comprised of both plants and animals in similar proportions), insectivore (>50% of arthropods in diet) and gummivore (diet dominated by plant exudates). We include in the trophic guild data base the percentage of fruit, leaves, flowers, seeds, animal matter, nectar and other, when available. We also include, when available, the study region, data type (e.g., feeding records, feeding time), study duration, group size and other general comments (e.g., when fruits and seeds are pooled). Body mass is expressed in kilograms; values for this variable can represent reported individual values, reported averages, or calculated averages (when a study included more than one body mass datum). When available, we separately report the mean body mass of adult males and females. Regarding habitat type, we include seven categories of major natural habitats in which a species occurs: (1) forest, includes ecosystems such as tropical wet forest, cloud forest, dry forest, montane forest, temperate forest and semideciduous forest; (2) savannah, includes savannah forest and savannah mosaics; (3) shrubland, includes ecosystems dominated by shrubs, such as scrub, brush and bush; (4) grassland, includes ecosystems mainly composed of grasses and other herbaceous plants; (5) wetlands, includes ecosystems such as swamps, flooded forest, swampy forest, wetlands and mangroves; (6) rocky areas, includes ecosystems such as inland cliffs and mountain peaks; and, (7)  www.nature.com/scientificdata www.nature.com/scientificdata/ geographic realms: Asia, Mainland Africa, Madagascar and Neotropics. African primates were classified in two groups, Mainland Africa and Madagascar, because these two land masses span the distribution of two highly divergent primate suborders (catarrhines and strepsirrhines, respectively). These methods are expanded versions of descriptions used in our recently published study 10 . Data files are stored in Zenodo 23 .

Data Records
The complete database for the ecological traits of primates consists of seven different data files with descriptive names (Table 1). Data files are stored in Zenodo 23 . We also include a text file "References.txt" that contains all the references included as numbers in the data files. The first row of each data file is the header containing the variables' names; each of the following rows presents data for a single primate species and a single information source. A given species can appear in more than one row because in some cases we included ecological traits from more than one source.

technical Validation
Most of the records included in the database are based on published material in peer-reviewed scientific journals and books, and thus we have confidence in their accuracy. Also, for each specific datum we include the corresponding reference in the database, allowing users to both assess the validity and consult the original sources. Moreover, we have carefully checked the database for possible redundancies and errors. When a specific datum was considered non-reliable (e.g., very extreme or contradictory values, and values obtained with questionable methodology) we did not include it in the database. We also used the 'validate' 24 R package to check the database for structural integrity (i.e., its internal organization). Our aim is to keep updating data for the traits recorded and incorporate other traits such as social attributes and other physical measures in the future. Data will be corrected and updated if any errors or updates are reported to the corresponding author.

Usage Notes
We would appreciate if researchers cite the database stored in Zenodo 23 in the specific version used, as well as this publication, when using all or part of the database.

Code availability
Code for the technical validation can be found archived in the Zenodo repository as "Test_validation" 23 . www.nature.com/scientificdata www.nature.com/scientificdata/ acknowledgements We gratefully acknowledge the Subject Editor (Karen Strier) and two anonymous reviewers for their detailed reviews of the database and valuable suggestions on the manuscript. CONACyT (Grant 2015-253946) provided financial support to V.A.-R., R.A.-G. and C.G.-A., C.G.-A. and R.A.-G. received a scholarship from CONACyT and C.G.-A. received financial support from Rufford Small Grants (18689-1). We are also grateful for the support (infrastructure, logistics and administration team) provided by the Instituto de Investigaciones en Ecosistemas y Sustentabilidad, UNAM. Part of the data compilation was done while C.G.-A. was on a research visit at the Imperial College London, UK.