Research Article |
Corresponding author: Rodrigo Antônio Castro-Souza ( rodrigodesouzaac@gmail.com ) Corresponding author: Joaquín Hortal ( jhortal@mncn.csic.es ) Academic editor: Mark John Costello
© 2025 Rodrigo Antônio Castro-Souza, Juliana Stropp, Luiz Felipe Moretti Iniesta, Richard J. Ladle, Neucir Szinwelski, Geiziane Tessarolo, José Alexandre Diniz-Filho, Thadeu Sobral-Souza, Joaquín Hortal.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Castro-Souza RA, Stropp J, Iniesta LFM, Ladle RJ, Szinwelski N, Tessarolo G, Diniz-Filho JA, Sobral-Souza T, Hortal J (2025) Mapping the status of global taxonomic knowledge of Orthoptera (Arthropoda, Insecta). Frontiers of Biogeography 18: e145455. https://doi.org/10.21425/fob.18.145455
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The status of taxonomic knowledge varies across the Globe. Quantifying and mapping the geographic patterns of taxonomic status is essential to prioritise regions that require greater attention from the taxonomic community. Here, we compiled all valid orthopteran species names and their synonyms, extracted from the Catalogue of Life (CoL) and allocated them geographically, based on data from the Global Biodiversity Information Facility (GBIF) and the Orthoptera Species File (OSF). This allowed us to create measures of taxonomic effort, based on the date of species descriptions and the number of associated synonyms and combine them across space. Our analyses show that the descriptions of currently valid species increased exponentially since the 19th century, with a temporary decline following World War II, while synonyms outpaced the number of valid species until the 1980s. The number of taxonomists also increased over time, with declines after World Wars, followed by a significant rise from the 1950s onwards, continuing through the 21st century (with > 100 taxonomists currently active). Per-taxonomist description rates transitioned from highly variable before the 20th century to consistent rates of 5–10 species annually with collaborative efforts. Tropical regions and the Southern Hemisphere hold the majority of recently described species names with fewer associated synonyms, indicating a predominance of alpha taxonomy and highlighting the need for greater taxonomic efforts. In contrast, temperate regions, particularly in Europe and south-western Asia, contain the majority of older names and synonyms, indicating a predominance of beta taxonomy and regions that have been more thoroughly studied. Our findings are discussed in the context of sociopolitical factors, scientific investments and the history of taxonomy. Finally, we propose a framework that makes the links between taxonomy and macroecology accessible for biodiversity in the era of Big Data.
Alpha taxonomy (i.e. the description of new species) and Beta taxonomy (i.e. the revision of the taxonomic status and relationships of already described taxa) vary across space and time.
We present a framework that connects taxonomy and macroecology, allowing us to assess taxonomic trends to provide information for biodiversity studies in the era of Big Data.
We mapped alpha taxonomy rates using the date of species descriptions across space and beta taxonomy rates using the number of synonyms associated with each species.
We combined alpha and beta taxonomy rates to understand the global taxonomic status of Orthoptera (Arthropoda, Insecta), the sixth most species-rich insect order, which includes grasshoppers, crickets, katydids and relatives.
In tropical regions, orthopteran taxonomy is recent and has few revisions, while in temperate regions, it is older and more consolidated.
Alpha taxonomy, beta taxonomy, biodiversity knowledge shortfalls, macroecology, Orthoptera, taxonomic uncertainty
Taxonomy is a theoretical and practical science that deals with biodiversity classification (
The discovery and naming of new species represents the first stage of taxonomy, known as alpha taxonomy (see
The second stage of taxonomy, known as beta taxonomy, involves understanding of relationships amongst already described species and higher taxa through their systematic revision (see
Mapping alpha and beta taxonomy across space can help unveil the mechanisms that have driven or limited taxonomic development in different regions, as well as highlight areas with varying levels of taxonomic stability. This, in turn, can enable the prioritisation of locations and territories that need more taxonomic attention in the form of new species descriptions, taxonomic revisions or both. However, to date, very few broad-scale investigations into these issues have been conducted, possibly due to a lack of a strong connection between taxonomic practice and macroecological research (see
Here, we combine the date of description of both valid species and synonyms and the number of synonyms associated with each valid species name, as proxies for variations in alpha and beta taxonomy across space, in order to map the global trends in the status of taxonomic knowledge. We hypothesise that beta taxonomy is predominant in Europe and temperate regions of the Northern Hemisphere, as the history of taxonomy began with European naturalists in the 18th century. The early description of a large portion of species from these regions should have allowed a subsequent focus on taxonomic revisions. Moreover, the higher research capacity (i.e. advanced technology, specialised expertise, funding and infrastructure) of many countries in the north temperate areas is predicted to be associated with a wider range of contemporary approaches to taxonomy, facilitating higher rates of taxonomic change. In contrast, we expect alpha taxonomy to be predominant in Tropical Regions, reflecting both the megadiversity yet to be documented in these regions, as well as a scarcity of taxonomists and technical capacity. These same factors should also lead to fewer taxonomic revisions.
We test the above hypotheses for the global diversity of Orthoptera (Arthropoda, Insecta), the sixth richest order of insects today, including crickets, grasshoppers, katydids and relatives. Since the publication of “Systema Naturae” (Linnaeus 1758), where only 59 orthopteran species were described for science, the taxonomy of this group has captivated a large community of orthopterists worldwide, resulting in over 29,000 valid species currently described (
We built a checklist of valid species names (SN) and associated synonyms (SYN) using the taxonomic classification organised from the Orthoptera Species File (OSF). This classification is provided in Darwin Core at the Catalogue of Life (CoL), where it contributes to the higher classification (Suppl. material
Filtering process and identification of valid species names and synonyms in the Catalogue of Life (CoL) considering all taxonomic components (genus + specific epithet + author(s) + year). The first step consisted in removing fossils, subspecies and synonyms of subspecies (Tax. status). In the second filter, we counted proposed names until 2023 (Tax. timeframe). Finally, the third filter consisted of removing duplicate records in valid species names and synonyms, disregarding synonyms resulting from subgenus syntax (Tax. ambiguities).
Based on the filtered data, we built temporal species accumulation curves, which describe the accumulation of species names over time and the quantity of new names proposed per year (see
Using the valid species names filtered in CoL (Fig.
We calculated the number of valid species names within grid cells (pi) of 2° resolution (using Projected Coordinate System EPSG-4326). This macroscale approach reduces omission errors in the distribution of many species (see
Workflow for building a taxonomic knowledge status map: (A) build a checklist of Orthoptera names (i.e. valid species names and associated synonyms) using data from the Catalogue of Life (CoL); (B) access the geographic distribution of the number of valid and total species names integrating data from the Orthoptera Species File Online (OSF) and the Global Biodiversity Information Facility (GBIF); (C) calculate two indices representing alpha and beta taxonomy: alpha temporal (AT) and beta proportion (BP); (D) build maps of alpha and beta taxonomy, rasterising the indices of AT and BP in each 2° grid cell with data on orthoptera species; and (E) combine the AT (y axis) and BP (x axis) indices into a bivariate taxonomic knowledge status map.
We calculated two different indices as proxies for alpha and beta taxonomy: Alpha Temporal (AT) and Beta Proportion (BP) for all cells of our 2° global grid. Both indices were based on the spatial distribution of currently valid species names obtained from their records, rasterised and mapped at the same 2° grid scale (Fig.
The alpha temporal index measures the temporal distribution of species-naming efforts at different cells. AT assumes that cells with recent species names indicate ongoing taxonomic efforts, while cells with older species names do reflect efforts made in the past or the absence of new efforts – in the case that no recent names are present in the cell. To calculate this index, we fitted the year of description of each valid species name to an exponential decay curve, derived from a polynomial function of degree 4, considering the time span of all names. This curve provides higher weights to older species names; as taxonomy advances with time, the weight for descriptions decreases exponentially. Specifically, we assigned an alpha temporal value for each valid species name ATsn (Eq. 1), based on the time elapsed from the most recent authorship year to the oldest (2023 to 1758 in our data), where species described in 1800 have values of ca. 0.5 and species described in 1900 ca. 0.1 (Fig.
AT sn = (ti − tmax/h)4, Eq. 1
where ti is the year of authorship of the investigated valid species name, tmax is the year of the most recent authorship in the checklist of the valid species names and h is the maximum value of the difference between the years when the names were described. Subsequently, taking the values previously assigned to each valid species name (ATsn), we calculate the average value of alpha temporal for each location (pi), based on the geographical occurrence of each species (Eq. 2).
AT pi = μATsn(pi) Eq. 2.
AT values range from ~ 0 to 1; we assume that when AT ~ 0 or ≤ 0.25, it indicates that the majority of valid species names in a cell correspond to recent descriptions, meaning alpha taxonomy is predominant. Conversely, values close to ≥ 0.75 or 1 indicate that a large proportion of valid species names in a cell are old, meaning that alpha taxonomy is currently not predominant (Fig.
The Beta Proportion index measures the portion of synonymised species names in a given location. We assume that the most studied sites would have more synonyms per species because of the greater effort for thorough taxonomic revisions, although this is not a general rule in taxonomy because of the priority rule in the Zoological Code of Nomenclature (see
For each cell (pi), we calculated Beta Proportion using Eq. 3:
BP = 1 − [sn(pi)] / [tn(pi)], Eq. 3
where BPpi is the result of 1 minus the ratio of the number of valid species name occurrences (sn) to the total number of name occurrences (tn) for each location (pi) and tn is the result of sn plus all associated synonyms for a given location (pi) (
BP values range from ~ 0 to 1; BP ~ 0 indicates a low proportion of synonyms relative to the existing species at the location, suggesting that beta taxonomy is not predominant. In contrast, values close to 1 reflect a high proportion of synonyms, indicating that beta taxonomy is predominant. It is important to note that it is not possible to determine whether beta taxonomy is recent or older (Fig.
To identify areas with different levels of alpha and beta taxonomy, we combined the two alpha temporal and beta proportion indices into a bivariate map. For this, we divided the AT and BP values (X and Y axes, respectively) into octiles (eight quantiles) and assigned different colours to different combinations (see
The accumulation of valid species names of Orthoptera has increased exponentially since the mid-19th century, with a brief decline just after World War II (Fig.
Considering the ratio of number of species names described by the number of taxonomist over time, the highest fluctuations occurred before the 20th century (with cases when a single taxonomist could describe more than 50 species); while more recently, each active taxonomist tends to describe between 5 and 10 species per year (Fig.
Our dataset included 76% of currently valid orthopteran species names. Considering only valid species names with synonyms (i.e. SYN ⊆ SN), we obtained information for 8,080 out of the 10,502 valid species names in the Catalogue of Life, representing 77% of all currently recognised species. We obtained spatial information for 21,945 of the 28,928 orthopteran valid species names recorded in CoL. Orthoptera valid species records cover all terrestrial regions of the world, except for northern Africa, western China and the Polar Regions (Fig.
The spatial distribution of the total number of names follows the same pattern (Fig.
Geographic distribution of the number of valid species names (A) and the total number of Orthoptera names (B). In (A), dark red colours indicate higher richness of valid species names, while light beige regions indicate lower richness. In (B), dark regions present higher total numbers of names (i.e. number of valid species names + number of associated synonymous names), while light colours indicate a lower total number of names.
Based on the Alpha Temporal index, the most recently described valid species (i.e. AT ≤ 0.5) are primarily found in tropical latitudes, whereas older species (i.e. AT ≥ 0.75) are mostly located in temperate latitudes, particularly in Europe and western Asia (Fig.
Spatial distribution of Alpha Temporal (A) and Beta Proportion (B) indices for Orthoptera names. In (A), yellow and green indicate older species names, blue represents more recent species names. In (B), yellow and green indicate a higher number of synonyms associated with valid species names, while blue corresponds to regions with lower proportions of synonyms.
The bivariate map of alpha and beta taxonomy reveals at least four main types of taxonomic trends around the world (Fig.
Interestingly, many regions have received intense taxonomic work in the last decades, coupling species descriptions with taxonomic reviews, characterised by recent species and many synonyms (i.e. low AT and high BP, blue in Fig.
Taxonomic knowledge status map resulting from the combination of alpha temporal (AT, X axis) and beta proportion (BP, Y axis). Regions in blue indicate high numbers of recently described species and synonyms; regions in purple indicate older species names together with many synonyms; regions in grey indicate recent species and few synonyms; and regions in yellow indicate older species and few synonyms.
Our analyses show that the number of newly-described Orthoptera species has increased exponentially over time (although this varies depending on the time frame, such as 1-, 5- or 10-year intervals), indicating that we are still far from reaching a near complete knowledge of the global diversity of this order. However, also importantly, the decline in the number of recorded synonyms over time may reflect a maturation of taxonomy, compared to previous decades. These results reinforce the essential and irreplaceable role of taxonomists, highlighting the need to address new species discoveries, but also that it is crucial that they go hand-in-hand with taxonomic revisions, especially considering the variation across taxonomic groups and the spatial heterogeneity of these patterns, as well as the adoption of new characters for species delimitation (see below).
Since 2014, the number of taxonomists describing new species actively working on Orthoptera has surpassed the mark of 100 (counting all taxonomist authors by valid species name). Currently, each taxonomist describes, on average, between 5 and 10 species per year, often in collaboration with other taxonomists. Although these figures may seem modest, it has driven the description of 200 to 400 new species per year. These findings indicate that Orthopterology is becoming an increasingly collaborative science and that training new taxonomists is essential if we are to catalogue the majority of species before they possibly become extinct in the Anthropocene. Therefore, maintaining this number of active specialists and an active collaboration network amongst them could be strategic goals for ‘The Orthopterists’ Society’, which, in addition to meetings, may be achieved through international specialisation courses that train taxonomists from different parts of the world.
The accumulation of valid species names after World War II does not apply uniformly across all groups within this order. For instance, both the number of taxonomists and the rate of species descriptions have declined since the 20th century for Acridomorpha, which comprises approximately 9,000 species (about one-third of all known Orthoptera species) (
The accumulation of synonyms and the proportion of synonyms described per year have declined, particularly since the second half of the 20th century. This trend likely stems from two opposing factors, which can influence the systematics of Orthoptera in different ways. On the one hand, certain regions (e.g. Europe and Southeast Asia) appear to have reached a consolidated knowledge as extensive phylogenetic studies and taxonomic revisions have been undertaken in recent decades which will have corrected many synonyms. On the other hand, in regions where taxonomy, arguably, remains in its early stages (i.e. grey areas), the decline may indicate a deficiency in synonymy detection, despite the ongoing description of new species. This limitation may introduce an additional source of uncertainty in biodiversity studies (
One significant factor influencing global taxonomy in recent decades is the availability and accessibility of online databases. Their benefits include maximising access to species descriptions, taxa distributions, data infrastructures and community discussions (
The increasing number of Orthoptera taxonomists over time results from various social and traditional factors. In some cases, it reflects the continuity of established taxonomic schools, where senior taxonomists often train networks of new taxonomists in specific taxa. Collaborative practice in taxonomy is also relevant (see
The taxonomic status varies significantly across the world, depending on the taxa (
For Orthoptera, the tropical regions contain the majority of recently described valid species, with fewer associated synonyms. This supports the previous hypothesis that these regions still require extensive taxonomic revisionary efforts (i.e. beta taxonomy), given the continued under-representation of their estimated species richness. On the other hand, temperate regions are dominated by older valid species names and a higher proportion of synonyms, reinforcing the idea that beta taxonomy is more prevalent in these areas.
The widespread distribution of areas with recent names and few synonyms reflects an emerging orthopteran alpha taxonomy, but little beta taxonomy, characterised by species discoveries which are not yet accompanied by comparing taxa across regions and refining their classifications through taxonomic revisions. This can be attributed to a historical shortage of taxonomists in megadiverse regions – the taxonomic impediment sensu
This contrasts with the situation in the historical centres of taxonomic research, as in Europe or North America, for instance, which are currently more active on beta taxonomy. These areas already hold a long history of taxonomic efforts, with a large number of taxonomic changes that result in a combination of older valid names and many synonyms. It is most likely that these regions are taxonomically saturated to some extent, with a significant proportion of their actual species richness already known, described and included in taxonomic reviews or catalogues (see
A large number of areas hold, on average, recent names and numerous synonyms. In these regions, the active research in both alpha and beta taxonomy couples the discovery of new species with taxonomic reviews and the publication of synonym lists. In some regions, such as in North America, these areas are adjacent to territories with older names and a large proportions of synonyms. This may indicate that, although regional checklists were previously thought to be complete (e.g.
Very few areas hold a combination of older names and few synonyms, which indicates only past inventory effort, but little or no revisionary work. This may be due to two distinct, yet non-complementary, phenomena. In the best-case scenario, in these areas, current taxonomic knowledge comes from studies that produced classifications well-structured in terms of nomenclature and taxonomic relationships in the past, which have persisted until the present day, even with evidence coming from new taxonomic approaches. Alternatively, the current regional checklists in these regions come from taxonomic work conducted in the past that has not been revisited or revised later on. Importantly, a significant part of these areas is distributed in countries where socio-political impediments have occurred in the past or are occurring nowadays (
It is worth noting that the global distribution of taxonomic knowledge depicted by our work is not only the result of taxonomic work, but also of sampling effort. Regions with frequent field surveys tend to have more species recorded in their checklists, leading to more opportunities for taxonomic effort, such as the identification of new species or the revision of already described ones. Conversely, areas with historically low levels of sampling effort can appear under-represented in terms of the occurrence of newer species simply because fewer surveys have been conducted there, not because they have fewer species. The regions with continuous taxonomic effort often have a history of consistent sampling and research investment, leading to more refined taxonomic studies (
This relationship between sampling effort and taxonomic progress is further reflected in how species names are distributed globally. The indices of alpha and beta taxonomy rely on the spatial distribution of valid species names around the world. However, most species distribution data are limited to fieldwork observations and digitised records from natural history museums. In general, these limitations tend to bias species occurrences toward roads, navigable rivers and major cities, leading to spatially skewed sampling efforts (
In this context, it is not surprising that regions with a higher concentration of natural history museums tend to have a lower proportion of undescribed species (Linnean shortfall) and better-documented occurrences of already described species (Wallacean shortfall). Supporting this inference, the largest natural history museums are located in North America and Europe. Indeed, most museums that house more than 1 billion objects (e.g. natural and cultural collections) are found in either the USA or central and northern Europe (
Most of America was the target of expeditions by European naturalists focused on inventorying natural resources and, amongst them, species (
Based on our results, this latitudinal gradient also reflects in well-known regions holding revisions (old AT and high BP), indicating the influence of, for instance, European expeditions in America over time on the representation of fauna. These expeditions typically landed on the eastern coast of the Americas and moved inland, documenting the biodiversity of the New World (
Even though taxonomy has played a pivotal role in biological science since the 18th century proposal of Linnaeus’ Systema Naturae, assessing the current status of taxonomic knowledge for any taxon remains crucial. It helps identifying understudied and undersampled areas requiring further attention and anticipates future trends in systematics, especially in relation to related fields such as molecular phylogenetics, biogeography and ecology. In general, spatial biases in both alpha and beta taxonomy can be influenced by various factors, including a lack of funding for monitoring and biodiversity studies and the shortage of trained taxonomists. Importantly, taxonomic advancements have largely been driven by increased knowledge from regions with consolidated and emerging research groups, for Orthoptera and other arthropod taxa. This is evident in regions with high alpha temporal and beta proportion, which are known for having historical zoological collections, curatorial assistance and scientific funding. In this sense, species richness revealed through taxonomic studies is not only linked to biodiversity patterns, but also driven by social, cultural and political factors, such as governmental funding, logistical challenges, the exodus of type material from its original locality and the shifting priorities of different historical periods.
We show that mapping the status of taxonomic knowledge allows the identification of regional trends in species description and revision. For the order Orthoptera, our model highlights that, in South America, Africa, Southeast Asia and Australasia, taxonomic efforts are needed not only for species description, but also for reviewing and integrating phylogenetic and biogeographic studies. Bearing this in mind, in the short term, tropical zones in grey and yellow in our map should prioritise alpha taxonomy, as these areas have a large species debt. In the long term, however, these same regions will need to advance towards the development of beta taxonomy, allowing for the revision and refinement of existing taxonomic knowledge. Additionally, we suggest that, in the short term, tropical zones in purple should be revisited by contemporary taxonomists, as these areas have great potential for the discovery of new species, especially through integrative approaches.
With this work, we aim to provide a foundation for exploring future challenges in Orthopterology, for both established taxonomists and new trainees, as well as for ecologists and biogeographers. However, more importantly, the framework presented here brings together taxonomy and macroecology in an accessible way for biodiversity studies in the era of Big Data.
We are indebted to the developers of Catalogue of Life (CoL), Global Biodiversity Information (GBIF) and Orthoptera Species File (OSF) that provided digitally accessible data. We are also grateful to the team at the Laboratory of Macroecology and Biodiversity Conservation (MacrEco) of the Federal University of Mato Grosso (UFMT) and the National Institute of Science & Technology (INCT) in Ecology, Evolution and Biodiversity Conservation (EECBio), based at the Federal University of Goiás (UFG), for the valuable discussions that contributed to the development of this study. We also thank Mark John Costello, Libin Ma and Maria-Marta Cigliano for their highly valuable suggestions and reviews. RACS was funded by Coordenação do Aperfeiçoamento do Pessoal do Ensino Superior (CAPES – finance code 001), Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) and Centro Nacional de Pesquisa e Conservação de Cavernas (CECAV) and currently is funded by CAPES (grant PIPD 88887.107568/2025-00). NS was supported by the Araucária Foundation (n° 09/2021). TSS expresses thanks to FAPEMAT (project FAPEMAT-PRO.000274/2023) and CAPES (grant 88887.964855/2024-00). GT was supported by the National Council for Scientific and Technological Development (MCTI/CNPQ/CAPES/FAPS N° 16/2014 - PROGRAMA INCT). JH was supported by project NICED, grant PID2022-140985NB-C21 funded by MCIN/AEI/ 10.13039/501100011033 / FEDER, EU.
R.A.C.S., J.S., T.S.S. and J.H. conceived the study with input from all authors; investigation, all authors; data curation, R.A.C.S., T.S.S. and J.H.; formal analysis, R.A.C.S., T.S.S. and J.H.; visualisation, R.A.C.S. and J.H.; funding acquisition, J.A.D.F., T.S.S. and J.H.; writing – original draft, R.A.C.S., T.S.S., L.I. and J.H.; writing – review and editing, J.S., L.I., R.J.L., G.T., N.S. and J.A.D.F.; All authors have read, discussed the results and approved the final manuscript.
The datasets used in this study are available in the supplementary material. The most updated versions of the original data can also be downloaded from Catalogue of Life (CoL), Orthoptera Species File (OSF) and Global Biodiversity and Information (GBIF). The underlying code for this study is not publicly available, but may be made available to qualified researchers on reasonable request from the corresponding author.
Orthoptera data available in Catalogue of Life (table S1) (.tsv)
Fossil Orthoptera data available in Catalogue of Life (table S2) (.xlsx)
Orthoptera occurrence available in Orthoptera Species File (table S3) (.csv)
Orthoptera occurrence available in Global Biodiversity Information Facility (table S4) (.7z)
Orthoptera occurrence filtered in Orthoptera Species File (table S5) (.zip)
Orthoptera occurrence filtered in Global Biodiversity Information Facility (table S6) (.zip)
figure S1: Sensitivity analysis between synonyms and valid species names; figure S2: Density curves of alpha temporal and beta proportion indices; figure S3: Correlation between the number of valid species names and the total number of names; figure S4: Distribution of global scientific collections and types of Orthoptera; appendix S1: The ‘nomenclatural’ lumping; appendix S2: Assessing the sensitivity of alpha temporal and beta proportion indices; appendix S3: Distribution of global scientific collections and types of orthoptera (.docx)