Addressing biodiversity knowledge shortfalls in New World Helicopsychidae (Insecta, Trichoptera): Potential distribution, environmental gradients, and identification of conservation and research priority areas

acknowledge that our data are limited by current knowledge of species distribution, and future research may expand the distribution range of species. Distribution patterns and potential distribution in New World The initial step in addressing the challenges posed by BKS is the compilation of inventories, checklists, descriptions of new species, and new distribution records [36]. However, it is equally important to consider other aspects, such as ecology, biology, phenology, and ethology, which can inform biodiversity conservation policies and the consequences of anthropogenic interventions (e.g., climate change, suppression of riparian vegetation) [23,25,26,71]. This study estab- lishes the environmental gradients for the H. (Feropsyche) and H. (Cochliopsyche) species. Despite the considerable range of elevation, temperature, and precipitation observed across both subgenera, the data indicated a higher prev- alence of H. (Feropsyche) species in coastal areas of tropical and temperate environments, across a wide elevation gradient (0–5042 m a.s.l.), in low order streams and in areas with high rainfall (Figs 3 and 4). On the other hand, the H. (Cochliopsyche) species are restricted to tropical environments, with prevalence in rainforests and lowland areas. Of the distributional records, 55% are in higher order streams (>4th order) (Figs 3 and 4). The results presented here concur with previous hypotheses proposed by Monson et al. [72], Flint [62,63], and Johan-

son [41,45]. However, this study provides a comprehensive overview of the distribution and analyzes the differences in environmental space between the subgenera. In addition to the morphological differences, these factors contribute to the accumulation of divergence between the NW Helicopsychidae lineages. Currently, there is still no clear resolution regard- ing their phylogenetic relationship. Some hypotheses suggest that they are sister groups [73], while others posit that they are distant clades [30], further complicating their taxonomic placement. Despite its wide distribution in tropical regions, H. (Cochliopsyche) is largely restricted to the type locality or adjacent localities (S1 Table). Distribution records are scarce and generally consist of single collections with a small number of specimens (S1 Table). This is primarily due to the greater concentration of collection and research efforts on Trichoptera in small, oxygenated freshwater environments, where most of the group’s diversity is concentrated [34]. It is therefore rec- ommended that research efforts be directed towards the at inventorying of South America’s large rivers, with priority areas for H. (Cochliopsyche) studies (areas of high suitability and low research effort) including all South America’s large river basins (e.g., Amazon, Orinoco, Paraná, and São Francisco River basins), with a particular focus on the large size rivers (Fig 1C). The species of H. (Feropsyche) are distributed across a wide range of habitats, from the temperate areas of the

Nearctic Region to the Andean areas of Chile (Fig 1E). However, the areas with the highest environmental suitability are throughout Central America (including the Antilles), the Florida Keys, areas east of the Andes between Argentina and

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 18 / 22 Bolivia (Chaco domain), and the coastal region of northwest and east South America. Much of continental Central Amer- ica, the Antilles, and the northwest of South America have been well sampled (Fig 1D). However, areas of the Chaco domain, the northern portion and Central Corridor of Atlantic Forest, and high-altitude areas of the Cerrado and Caatinga domains remain poorly sampled (Fig 1E). These areas should be prioritized for research efforts, in agreement with the Brazilian caddisfly studies of Santos et al. [24], which indicated the to prioritize research efforts in the Northeastern and Central-Western regions of Brazil. The Patagonian subregion is one of the least researched in the NT, with both groups exhibiting low suitability for this region. However, 11 species of H. (Feropsyche) are found in areas with minimum temperatures below 0°C (e.g., H. (Fer- opsyche) borealis, −17°C; H. (Feropsyche) minuscula, −12°C) (Fig 1D). The data available do not permit us to determine whether the low suitability is (i) due to the absence of favorable environmental conditions, or (ii) to the scarcity of data, which resulted in the model’s limitation for this area. In general, there is a considerable range of distribution records for the Helicopsyche in the NW. The species are distributed across a wide environmental gradient, with H. (Feropsyche) being particularly notable for their ability to occur in various freshwater environments across the New World. This includes a wide range of altitudes,

temperatures, and precipitation levels, with a preference for low-order environments (Figs 1,3, and 4). In contrast, H. (Cochliopsyche) species typically occur in lowlands, environments with higher average temperatures, and a preference for large rivers, reflecting a restriction in their occurrence to large river basins (Figs 1,3, and 4). Thus, the results indicate that the two lineages have specialized in different niches: H. (Feropsyche) specialized in smaller (low-order) environments, while the H. (Cochliopsyche) specialized in larger (high-order) environments. Similar patterns are also observed in other Trichoptera lineages, including Hydrobiosidae (Amphichorema and Atopsyche), and Glossosomatidae (Culoptila and Cariboptila) [33]. The results of this study highlight critical conservation implications for Helicopsychidae species in the New World. The identification of environmental gradients and the discovery of species with restricted distribution in high-suitability and low research effort areas emphasize the need for targeted conservation efforts. Areas with high species richness and environ- mental suitability, such as parts of Central and South America, should be prioritized for further research and protection. The underexplored regions, especially tropical rainforests and the dry diagonal zones (Caatinga, Cerrado, and Chaco), represent significant gaps in knowledge and potential hotspots for biodiversity. Increasing faunistic surveys in these regions will not

only enhance species descriptions but also help mitigate the risk of extinction for species with narrow distribution ranges, which are more susceptible to threats such as climate change and habitat loss. Moreover, the conservation of riparian habitats, particularly in large river basins, is critical to maintaining the integrity of ecosystems supporting these specialized species. By addressing both Linnean, Wallacean, Haeckelian, and Hutchinsonian and other shortfalls, future research can refine conservation strategies and promote the preservation of these important freshwater ecosystems. Conclusion The results highlight recent progress in the taxonomy and distributional cataloguing of New World Helicopsychidae, partic- ularly within underexplored regions of the Neotropics. Despite a substantial increase in described species, significant gaps in biodiversity knowledge remain. Current estimates suggest that up to 75% of Helicopsyche species in the region are still undescribed, emphasizing the urgent need for continued research efforts—ideally incorporating all semaphoronts—to fully document and understand the diversity of NW Helicopsychidae The distribution of Helicopsyche species reveals complex patterns influenced by environmental factors such as ele- vation, temperature, and precipitation. Species of H. (Cochliopsyche) are typically associated with larger river systems in warmer climates and extensive river basins, whereas H. (Feropsyche) species show broader ecological tolerance,

occurring across diverse freshwater habitats throughout the Americas, particularly in low-order streams. These contrasting patterns reflect distinct ecological specializations between the two subgenera.

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 19 / 22 Sampling efforts across the New World remain uneven, with the majority of data derived from historically surveyed regions (e.g., the Antilles) or areas with established research institutions (e.g., Central America, the Amazon, and the Atlantic Forest). These regions, due to their high species richness, should be prioritized in conservation planning, especially those identified as biodiversity hotspots. However, vast areas remain poorly explored, including the Amazon lowlands, northern and central Atlantic Forest, high-altitude environments in the Caatinga and Cerrado domains, and tran- sitional zones between major phytogeographic domains. Targeted research in these underrepresented areas is essential. This study represents a foundational step toward a more comprehensive database of Trichoptera diversity in the New World. It provides critical tools for delineating biogeographic patterns and identifying knowledge gaps, thereby enhancing our understanding of Helicopsychidae diversity. The compilation presented herein establishes a valuable baseline for future taxonomic, ecological, and conservation-oriented studies on this family. and may inspire future research proposals for this family or other Trichoptera taxa using similar approaches. Supporting information S1 Table. Summary of georeferenced distribution data for all New World Helicopsychidae species. (XLSX) S2 Table. Summary of distribution in Freshwater Ecoregion (sensu Abell et al. 2015) and limits data for biogeocli-

matic variables of all New World Helicopsychidae species. (XLSX) S3 Table. Pearson correlation between environmental variables. (XLSX) S1 Fig. Boxplot with the result of Pearson correlation between environmental variables. (TIFF) S2 Fig. Boxplot with result of evaluation of distribution modelling algorithms using the area under curve (AUC) method. (TIF) Acknowledgments We extend our sincere thanks to Dr. Neusa Hamada, Dr. Ana Maria Pês, and Dr. Gleison R. Desidério for kindly providing data and permitting the analysis of material from the Laboratório de Citotaxonomia e Insetos Aquáticos of the Instituto Nacional de Pesquisas da Amazônia (LACIA-INPA). We are also grateful to the Museum of Comparative Zoology (MCZ) at Harvard and to Dr. Crystal A. Maier for granting access to, and assisting with the analysis of, biological material from the MCZ collection. Our appreciation further extends to Dr. Ralph Holzenthal and Dr. Robin E. Thomson at the University of Minnesota, and to Dr. Dikow Torsten of the Smithsonian Institution for kindly loaning specimens from institutional collec- tions for analysis at the MCZ. Author contributions Conceptualization: Rafael Pereira, Adolfo Calor. Data curation: Rafael Pereira. Formal analysis: Rafael Pereira. Funding acquisition: Adolfo Calor.

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