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. 2025 May 22;12:840. doi: 10.1038/s41597-025-05089-3

CheloniansTraits: a comprehensive trait database of global turtles and tortoises

Jiang Wang 1, Yunhao Xu 1, Huiwen Zhu 1, Chuanwu Chen 1, Yifan Zhao 1, Yanping Wang 1,✉
PMCID: PMC12098739  PMID: 40404775

Abstract

Turtles and tortoises (chelonians) possess a variety of ecological characteristics, including long lifespans and protective shells, which have enabled them to survive and adapt to environmental challenges since the Triassic period. However, many characteristics of chelonians have turned into disadvantages for their populations in the Anthropocene. Currently, there remains a lack of comprehensive data on the morphological, life-history, and ecological characteristics of all chelonians on a global scale. Consequently, our study aims to collect a complete trait database of global chelonians (CheloniansTraits), which may help bridge the knowledge gap regarding the identity and ecology of global chelonians and thereby aiding future conservation endeavors. We compiled 69 trait data for all 358 recognized chelonian species, utilizing ~2,000 literature sources, covering 33 morphological, 21 life-history, 7 ecological traits, and 8 conservation information. This database serves as a uniquely valuable resource for exploring evolutionary, biogeographical, and ecological inquiries related to chelonians, as well as elucidating key aspects of ecological strategy variation among species.

Subject terms: Biodiversity, Conservation biology, Biogeography

Background & Summary

Turtles and tortoises, or chelonians (Order: Testudines), tracing back to the Triassic period, have an older origin compared to other modern vertebrate clades1,2. Chelonians are comprised of more than 300 species that inhabit various aquatic and terrestrial ecosystems and are widespread worldwide, except Antarctica3,4. Over 200 million years, chelonians have developed a variety of unique morphological, life-history, and ecological traits that have enabled them to survive and adapt to environmental challenges5. They are also widely known for their ecological importance as mesopredators of animals and seed dispersers in ecosystems5,6.

In the Anthropocene, however, these unique traits have turned into disadvantages for Chelonian populations7–9. For instance, the shell, as an adaptive trait, provides shelter from the environment, shields against predators, enhances thermoregulation and serves as a valuable reservoir of fats, minerals, and water10,11. Nevertheless, the shelled-body restricts their mobility, especially terrestrial ones, making them easier to be captured and overexploited by humans12,13. Moreover, life-history traits such as long lifespan, delayed sexual maturity, and long generation times confer advantages for the survival of chelonians under long-term natural selection pressures1,14,15. Unfortunately, their long lifespan has also led many people to believe that consuming their meat and eggs or using them in traditional medicines will improve longevity16–18. Meanwhile, individuals with large body sizes and mature females are often sources of protein for local residents and the main targets of trade (e.g., in Amazonian cultures)9,19. Thus, the delayed sexual maturity and long generation times traits render them more difficult to recover from population losses9,17. In addition, chelonians have developed various abilities to adjust to extreme environmental conditions (e.g., thermal tolerance, resistance to food shortage, and anoxia tolerance), which make them withstand long-distance transportation in global trade20,21. In this context, populations of chelonians are declining rapidly and many species have gone extinct due to human activities4,8. For example, over half species of chelonians are threatened to extinction, as assessed by the International Union for Conservation of Nature (IUCN) Red List22. To date, chelonians rank among the most threatened vertebrate groups, second only to primates9,23,24.

Understanding the species traits of chelonians is vital for developing effective conservation strategies25–27. However, the database on the morphological, life-history, and ecological traits of chelonians is generally lacking compared to other vertebrate groups, such as mammals, birds, and amphibians28–32. In particular, the identification of chelonians largely relies on their morphological characteristics, while it remains a big challenge for customs and forestry law enforcement personnel regarding the complexities of these features. Despite that several turtle identification guides have been published, many of them lack effective identification guidance for non-experts to accurately judge and identify turtles. Furthermore, previous studies that compiled body size datasets, such as Itescu et al.33 and Regis and Meik34, have certain limitations. These limitations stem from insufficient representativeness due to a small number of species examined and traits included or the use of generalized size approximations rather than more precise size data. Aside from these compilations of turtle body size datasets, Colston et al.35 and Rodríguez-Caro et al.36 also published two datasets on the ecological or life-history traits of turtles. However, as summarized by Colston et al.35 and Rodríguez-Caro et al.36, these data are scarce and contain many missing values. While we acknowledge that past trait databases can be important and have laid the foundation for the availability of datasets, compiling a global dataset on chelonians will help drive the conservation of these species in regional or global contexts. The most recent trait database related to chelonians is ReptTraits37, but it only focused on the ecological traits of chelonians, lacking specificity and practicality for general use. Moreover, the taxonomic checklist of the ReptTraits37 is a little bit confusing, which includes subspecies and subpopulations of chelonians (e.g., Elseya caelatus, Chelodina ipudinapi, Elseya auramemoria). The ReptTraits also unnecessarily included some traits unsuitable for chelonians, such as venomous, venomous fangs, and reproductive mode37. In fact, after excluding these trait information, only eight (since maximum Longevity is also misclassified) morphological traits of chelonians were collected in the ReptTraits database. Finally, the ReptTraits dataset has poor data completeness on all the life-history traits (<40%), except for the body size (actually, the maximum body mass data primarily cited is converted data based on allometry sourced from Meiri et al.38) and the geographical distribution of chelonians32. Therefore, the ReptTraits dataset is inadequate to represent the full appearance of chelonians and also impractical for species conservation.

To bridge the gap, we collected a complete trait database of global chelonians (CheloniansTraits) that encompasses morphological traits, life-history traits, ecological traits, and conservation information. We made it possible by three-year efforts on collecting trait information from ~2,000 literature spanning from 1758 to 2024, related academic books such as “Turtles of the World: A Guide to Every Family”39 and “The Conservation Biology of Tortoises”40, existing morphological and life-history databases (e.g., Itescu et al.33, Regis & Meik34, Colston et al.35), the IUCN Red List (version 2024-2)22, the Reptile Database41. Our trait database was double-checked by specialists in this field to ensure completeness and accuracy. Overall, 358 currently recognized species (including the few that are known to have gone extinct in recent centuries) belonging to 14 families were checked with species-level identities. For these species, we collected 69 traits, including 33 morphological traits, 21 life-history traits, 7 ecological traits, and 8 variables related to conservation information. We aim to enrich our understanding of global chelonians through this trait dataset. We hope that our dataset can contribute to the development of studies and efforts on effective species conservation of chelonians.

Methods

Species checklist

We obtained the species-level taxonomy of chelonians based on the taxonomic checklist of the Turtle Taxonomy Working Group (357 species)42, the Reptile Database41, and a review of scientific literature sources spanning the period from 1758 to 2024. Some species that require further evidence for their species-level identity were categorized as controversial species. Finally, we included 358 chelonian species with confidently identified records in our study. For each species, we recorded the information on species order, family, genus, binomial species name, English name, and year of description from the Turtle Taxonomy Working Group42.

Data extraction procedure

To compile a comprehensive trait dataset for chelonians, based on the initial survey, we selected 33 morphological traits, 21 life-history traits, 7 ecological traits, and 8 conservation information related to chelonians. These trait data reflect a variety of the unique morphological, ecological strategies, and functional roles of chelonians. They are primarily derived from extensive classification literature, academic books, and life-history databases22,41–43. Our data collection process primarily adheres to the following four steps (Fig. 1): (1) First, we confirmed 358 turtles and tortoises at the species-level based on the taxonomic checklist of the Turtle Taxonomy Working Group42, the Reptile Database41 and scientific literature sources (from 1758 to 2024). (2) Second, we integrated data from six primary datasets (Itescu et al.33, Regis & Meik34, Colston et al.35, Xiao et al.44, Rodríguez-Caro et al.36, and Oskyrko et al.37), compiling the raw data. (3) For morphological, life-history, ecological traits, and conservation information, we supplemented them by consulting literature43,45,46 and using dedicated websites22,41,46. We then checked raw data and newly-discovered species and deleted errors. (4) Finally, we supplemented missing data by searching the keyword ‘species name’, ‘turtles’, ‘morphological’, ‘reproductive’, ‘diet’, ‘active pattern’, and ‘clutch size’, on the Reptile Database (http://www.reptile-database.org)41, Animal Diversity Web (https://animaldiversity.org)46, the IUCN Red List (https://www.iucnredlist.org)22, Web of Science (https://www.webofscience.com), and Google Scholar (https://scholar.google.com) and integrated the chelonians traits database. In cases of data conflicts, we collected various values including maximum, mean and range data for parameters such as body size, lifespan, clutch size, number of clutches per year, incubation period, and egg size as much as possible. However, the data of diet or activity time can vary due to life stages, observations or seasonal changes39,47,48. Therefore, we extracted these values for individual traits by choosing data for the adult stage or main active patterns in our study. For example, green turtles (Chelonia mydas) are omnivorous during some life stages, ranging from carnivorous on marine animals as juveniles to becoming mostly herbivorous on marine plants as adults39. We recorded adult green turtles eating sea grass and macroalgae in the detailed diet data, categorizing their diet as herbivorous. We divided the activity patterns of turtles according to the timing of key behaviors such as foraging, basking, nesting, and movement mentioned in the primary source. The activity patterns of Blanding’s turtles (Emys blandingii) were described as diurnal because rarely recorded Blanding’s turtles moving at night49.

Fig. 1.

Fig. 1

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Elucidation of the workflow for dataset creation of global turtles and tortoises.

Trait data

For the data of 16 body size traits, we collected the carapace and plastron length data separately for adults, males, females, juveniles, and unknown-gender individuals. The data of head length and width for adults, maximum mass, and hatchling/neonate mass are also aggregated (Fig. 2). When multiple resources are available for a species, we prioritized the resource that contains the most extensive and comprehensive feature descriptions (e.g., TTWG 202142). However, body size values are typically population-dependent, varying geographically, latitudinally, and altitudinally (e.g., Testudo hermanni)42,50. They are influenced by sample sizes and potentially biased collection methodologies42. We therefore also included ranges in the parameters to compare against and provide a comprehensive database. Non-continuous morphological traits, such as scutes, tentacles (pair), and number of toes/fingers in the forelimbs and hindlimbs, are often incompletely or inadequately documented in published literature or books. Therefore, we collected these data by referencing the descriptions of scales and scutes provided by Pritchard51 and using high-quality pictures from sources such as The Reptile Database41 and TTWG42. Specifically, approximately 90% of the non-continuous morphological trait measurements were derived from images provided by TTWG42.

Fig. 2.

Fig. 2

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Percentage of data completeness for each trait released in the database.

Our dataset also includes 21 life-history traits. Turtles and tortoises have diverse diets, foraging modes, and activity time37,39,49. The long lifespan of chelonians is a unique life-history trait9,39. In addition to collecting the recorded lifespan of each species (in years), we have compiled details on their maximum lifespan (in years) as well. The adult survival rate and juvenile survival rate, which are associated with survival, are calculated as the arithmetic mean of survival values for specific life stages36.

We also gathered the reproductive traits of chelonians from literature descriptions46,52,53, including age at sexual maturity, breeding season (in months), incubation period (in days), clutch size (reported as minimum, maximum, and average values), number of clutches per year, egg size, eggshell texture and shape, and sex-determination systems.

For ecological traits, such as range sizes (km2), microhabitat, habitat type, endemicity, biogeographic regions, distribution, and presumed Historic Indigenous Range (km2), we reported these traits based on numerous previous studies35,42,54.

For conservation information, we obtained the threat status, assessment criteria, population trends, threats, use, and trade from the IUCN22. The threat status of chelonians, as assessed by Rhodin et al.45, was also included. Trade records spanning the period from 1758 to 2023 were collected from the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES, https://trade.cites.org/)55, and the presence or absence of trade records (yes or no) for each species was also reported. Finally, we examined publicly available genetic data (GenBank, https://www.ncbi.nlm.nih.gov/)56 for each species and reported whether genetic data was accessible or not (yes or no).

Data Records

The dataset is open access and available at the Figshare repository as ‘CheloniansTraits dataset.xlsx'57. We created an Excel spreadsheet that includes both the dataset content and the column descriptions as separate worksheets (Table 1: CheloniansTraits data, Table 2: Main sources of trait value, Table 3: References, Table 4: Trait definitions and the completeness level, Table 5: Controversial species, Table 6: Type specimen, Table 7: Source of images)57. We provided the most comprehensive trait database of chelonians to date, encompassing 358 species from 14 families. In total, we extracted and collected the data from 2,000 literature sources and 20 databases. In addition to species order, family, genus, binomial species name, English name, and year of description, our dataset includes 69 traits. The trait completeness varied from 6.20% to 100%, with higher completeness of ecological traits and conservation variables, followed by morphological traits and life-history traits (Fig. 2).

Technical Validation

We implemented four steps to compile CheloniansTraits (Fig. 1) and meticulously reviewed the dataset to ensure consistency in variables. All data was manually gathered from the respective sources for each species and carefully recorded. To ensure traceability of the data, we incorporated both primary and secondary sources, clearly indicating their respective origins. Data sources that were difficult to interpret and lacked extractable raw data were excluded from the dataset. In addition, double control procedures were implemented to ensure the accuracy of the dataset. We confirmed and re-examined whether the problem data originated from transcription errors by cross-referencing the raw file with the source data. During the double-checking process, we verified the validity using multiple literature sources and made corrections or removed the data from the database as needed.

Usage Notes

Our database is the most comprehensive trait data available to date that has a full species coverage, detailed morphological traits for the first time, and a standardized referencing index. Notably, CheloniansTraits exhibited a high coverage of traits for each species, surpassing previous datasets for reptiles or chelonians. This database serves as a uniquely valuable resource for exploring evolutionary, biogeographical, and ecological inquiries related to chelonians, as well as elucidating key aspects of ecological strategy variation among species.

Acknowledgements

This study was supported by the National Natural Science Foundation of China (32271734 and 31971545). No permits for ethics approval or fieldwork were required for this project.

Author contributions

Y.W. conceived the ideas; J.W., C.C. and Y.Z. collected the data; Y.X. and H.Z. checked the data and added the primary sources of the data; J.W. led the writing of the manuscript with critical inputs from Y.W.; Y.W. supervised the work.

Code availability

No custom code was used to generate the described databases.w

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Citations

  1. NCBI Sequence Read Archivehttps://www.ncbi.nlm.nih.gov/ (2024).
  2. Wang, J. et al. CheloniansTraits: a comprehensive trait database of global turtles and tortoises, figshare, 10.6084/m9.figshare.28828241 (2025). [DOI] [PMC free article] [PubMed]

Data Availability Statement

No custom code was used to generate the described databases.w

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