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

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 1 / 22 OPEN ACCESS Citation: Pereira R, Calor A (2025) Addressing biodiversity knowledge shortfalls in New World Helicopsychidae (Insecta, Trichoptera): Potential distribution, environmental gradients, and identification of conservation and research priority areas. PLoS One 20(7): e0327580. https://doi.org/10.1371/journal.pone.0327580 Editor: Halil Ibrahimi, University of Prishtina, ALBANIA Received: March 18, 2025 Accepted: June 17, 2025 Published: July 28, 2025 Peer Review History: PLOS recognizes the benefits of transparency in the peer review process; therefore, we enable the publication of all of the content of peer review and author responses alongside final, published articles. The editorial history of this article is available here: https://doi.org/10.1371/journal. pone.0327580 Copyright: © 2025 Pereira, Calor . This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, RESEARCH ARTICLE Addressing biodiversity knowledge shortfalls in New World Helicopsychidae (Insecta, Trichoptera): Potential distribution, environmental gradients, and identification of conservation and research priority areas Rafael Pereira 1,2☯*, Adolfo Calor2☯ 1 Laboratório de Organismos Aquáticos – LOA, Programa de Pós-Graduação em Sistemas Aquáticos Tropicais, Universidade Estadual de Santa Cruz – (PPGSAT – UESC), Ilhéus, Bahia, Brazil, 2 Laboratório

de Entomologia Aquática – LEAq, Instituto de Biologia, Programa de Pós-Graduação em Biodiversidade e Evolução, Universidade Federal da Bahia (PPGBioEvo, UFBA), Salvador, Bahia, Brazil ☯ These authors contributed equally to this work. * rafaelpsilvape@gmail.com Abstract This study provides an integrative analysis of Helicopsychidae biodiversity in the New World (NW), examining their distribution patterns, environmental gradients, biodiver- sity hotspots, and biodiversity knowledge shortfalls. We estimated species richness, delineated potential distribution, and identified areas of priority for research/conser- vation efforts. Our estimates indicate that up to 75% of NW Helicopsychidae species remain undescribed, with a notable Linnean shortfall at the subgenus level: approxi- mately 50% of H. (Cochliopsyche) species and up to 77% of H. (Feropsyche) species are yet to be described. Knowledge of semaphoronts is also limited, with immature stages documented for only 15% of H. (Feropsyche) species and 6% of H. (Cochlio- psyche). Distributional records are concentrated in the Brazilian subregion (between 0°–24°S), and most species have a short environmental distribution gradient. Based on currently known, priority conservation areas are in the Antilles and the Tropical Forest. For H. (Feropsyche), priority areas include low-altitude and coastal regions with low-order streams, whereas H. (Cochliopsyche) conservation priorities lie in the large river basins, especially in the Amazon and Atlantic Forests. Future research

efforts should focus on H. (Feropsyche) in the Chaco and high-altitude areas of the Atlantic Forest, Cerrado, and Caatinga domains, as well as on H. (Cochliopsyche) across South America’s high order river (e.g., Amazon, Orinoco, Paraná, and São Francisco Rivers). Here, highlights recent advances in the taxonomy and distribution cataloguing of NW Helicopsychidae, despite significant progress, marjory of spe- cies remain undescribed, emphasizing the need for continued research. Although well-sampled regions like the Antilles, south Central America, Amazon coastal areas, and Central Atlantic Forest Ecological Corridor should be prioritized for conservation,

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 2 / 22 vast areas such as the Amazon lowlands, northeast and southwest Atlantic Forest, and transition areas of Atlantic Forest and Dry Diagonal remain underexplored. To the best of our knowledge, this study represents one of first empirical analysis of envi- ronmental gradients for Trichoptera species. It establishes a foundation for under- standing biogeographic patterns, environmental gradients, and the identification of biodiversity hotspots and potential distribution areas for Helicopsychidae in the New World. This work will guide future research and conservation efforts for Helicopsychi- dae and other Trichoptera groups in the region. Introduction The distributional patterns generally used the distribution of known species as a proxy [1], and, consequently, they are strongly related to biased collection efforts [2]. There are huge gaps in biodiversity knowledge related to areas neglected in terms of collection effort, fauna inventories, taxonomists, and investment in research, espe- cially in developing countries [3,4]. These factors cause collection bias and directly affect biodiversity knowledge [5–7]. In this context, some authors have defined the different knowledge gaps, or the Biodiversity Knowledge Shortfalls (BKS), in various categories such as gap of species knowledge is the Linnaean shortfall, the unknown species geographic distribution is the Wallacean shortfall, the gap of knowledge about semaphoronts of

species Haeckelian shortfall, the gap of knowledge about abiotic tolerance of species Hutchinsonian shortfall and unknown of evolution of groups is the Darwinian shortfall [8,9]. Among the tools available to face the biodiversity knowledge shortfalls, dis- tribution modelling can identify areas with a high probability of potential distribution (environmental suitability) [10]. The generated models can help to prioritize resources and efforts in areas with the probability of potential distribution of species [11]. On the other hand, anthropogenic actions have caused planetary-scale changes, which have caused the sixth mass extinction [12] with critical loss of biodiversity faster than our ability to catalogue it [13]. Among the ecosystems, freshwater environ- ments cover only 1% of Earth’s land surface but comprise around 10% of all species [14]. Levels of endemism in these environments are remarkably high, and about 20% of the New World (NW – here composed for West and East Nearctic, Neotropical and Patagonian region sensu de Moor & Ivanov) species are threatened sensu IUCN [15]. Due to these characteristics, freshwater environments have the most acute bio- diversity crisis among ecosystems [16]. The formulation of strategies based on large datasets is urgent to protect species, as well as the priority areas [17], and conse- quently reduce (or reverse) the decline of freshwater biota as much as possible [16]. Several insect orders have species with life stages occurring in freshwater ecosys-

tems, but some orders stand out as having primarily aquatic origins (e.g., Trichoptera) [18]. Among these amphibiotic orders, the caddisflies stand out as having the great- est richness and diversity of functional trophic groups [19]. Despite recent advances in caddisfly taxonomy and phylogeny, the order is underexplored and poorly known and reproduction in any medium, provided the original author and source are credited. Data availability statement: All relevant data are within the manuscript and its Supporting Information files. Funding: We thank the Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) for collecting permits. ARC acknowledge the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (under Grants 303623/2015-2). RP also thanks to the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES, finance code 001, PDS-CAPES-88882.453922/2019-01, and PRAPG-CAPES-88887.986811/2024-00) for the doctoral and post-doctoral fellow- ship. We thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), Programa de Apoio à Pós-Graduação (PROAP-CAPES) for financial support, and this study was financed in part by the CAPES – Finance Code 001 (PPG Biodiversidade e Evolução). We would like to thank Dra. Neusa Hamada, Dra. Ana Pes and Dr. Gleison Desidério for providing us with data from LACIA-INPA. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing interests: There is no conflict of interest.

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 3 / 22 in several aspects [e.g., 18–22]. Knowledge about Trichoptera species, distribution ranges, and biogeographic patterns is scarce, especially in the Neotropical region (NT) sensu de Moor & Ivanov [20,23–26]. Studies of Trichoptera have focused almost entirely on systematics, with few studies related to biogeography and a much smaller proportion concerned with understanding which factors limit the distribution of groups [25]. However, data related to the biology, environmental and biogeography limits are essential for understanding how species will behave both in scenarios of environmental impacts (e.g., deforestation, burning) and in scenarios of climate change [27]. Especially when it comes to taxa that play a key role in the trophic dynamics of freshwater and adjacent terrestrial ecosystems, such as the Trichoptera [19], and with factors such as temperature, rainfall, and riparian vegetation directly influencing the distribution of species [28]. Despite its importance, it has suffered alarming population declines, 2/3 (68%) of the caddisfly species experiencing population declines, 63% of the species threatened, and a local/regional extinction rate of 6.8% [29]. Among the caddisflies, Helicopsychidae von Siebold, 1856 are represented by two genera, the monotypic Rakiura McFarlane, 1973 (endemic to New Zealand), and Helicopsyche von Siebold, 1856 (circum-global distribution) [30]. Helicopsyche comprises 298 valid species in six subgenera: Cochliopsyche Müller, 1885, Feropsyche Johanson, 1998,

Galeopsyche Johanson, 1998, Helicopsyche, Petrotrichia Ulmer, 1910, Saetotricha Brauer, 1865 [31]. In the Neotropical (NT), Nearctic (NA) and Patagonian (PA) regions (sensu de Moor & Ivanov) have 146 valid species in two subgenera, H. (Cochliopsyche), with 17 Neotropical species, and H. (Feropsyche), with 129 species (including three fossil species from the Dominican Republic) widely distributed throughout the NW (NA with 13, NT with 125, and PA with two valid species) [31,32]. Helicopsyche is a genus recognized by larval cases built with sand grains, helically organized, and resembling snail shells [33]. Like most caddisflies, the immature stages live in freshwater ecosystems and, after completing the metamor- phosis, they emerge as winged adults associated with riparian forests [34]. Due to its wide distribution and occurrence in different freshwater environments of the NW [35], the genus represents a good biological model for initial studies that seek to identify and define distributional patterns of amphibiotic insect groups. Furthermore, understanding how these species are distributed and grouped can provide subsidies for the identification of under-sampled areas [36]. The subgenus Feropsyche was revised by Johanson (2002), and subsequent studies have primarily focused on spe- cies descriptions and new distribution records [e.g., 32,37–40]. In contrast, the subgenus Cochliopsyche has received comparatively less attention following its revision by Johanson (2003) [41], with most contributions limited to new distribu-

tion records [e.g., 42–44] and the description of a single new species, Helicopsyche (Cochliopsyche) nyurga Oláh & Oláh, 2022. The taxonomic literature on Helicopsyche also reveals a notable bias toward male specimens [33,45]. Species description based only on male adults is a quite frequent practice in caddisfly taxonomy and accurately sepa- rates species [24], as in some other insect orders [7]. For the most caddisflies, immature stages and females are unknown (an average of <15% of the immatures are known, and in 41 Neotropical genera, the immature stages are unknown) [46], demonstrating a knowledge shortfall related to other semaphoronts. This causes a genuine problem since most adult collections use light traps, which collect more adult females than males [47]. In addition, Helicopsyche species are mainly reported exclusively to type localities and/or adjacent localities (like the same district) [e.g., 32]. The combination of these two factors inevitably leads to knowledge shortfalls [8]. Understanding distribution patterns, environmental gradients, and biodiversity hotspots—along with identifying Biodiver- sity Knowledge Shortfalls (KBS) and prioritizing research efforts—provides crucial insights for addressing Wallacean and Hutchinsonian shortfalls and informs strategies for tackling other BKS. Additionally, this information forms a foundation for developing new research proposals, conservation actions, and enhancing understanding of biogeographic patterns

and the relationships between species and regions for New World Trichoptera. Within this framework, this study aimed to identify distribution patterns, establish environmental gradients, and model the distribution of Helicopsychidae subgenera in the New World.

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 4 / 22 Materials and methods Study area The New World is divided into three regions: the Nearctic, Neotropical, and Patagonian regions, with two transition zone: (i) Mexican transition zone comprise mountainous areas of central and southern Mexico and northern Central America, serves as a pathway connecting elements of the Nearctic and Neotropical fauna; and (ii) South American transition zone comprise highlands of the Andes between western Venezuela and northern Chile and central western Argentina [48]. Following these delimitations and using the distribution of Trichoptera as a proxy de Moor & Ivanov [20] divided Nearc- tic region into three regions: (i) West Nearctic (=Western subregion), (ii) East Nearctic (=Alleghany subregion), and (iii) Beringian “in part” (=Artic subregion) [14]. In addition to two regions covering part of Mexico, Central and South America, the Neotropical region (=Neotropical region) which includes Antillean, Brazilian, and Chacon subregions [sensu 48,49], and Patagonian region (=Andean or Chilean region) which includes Central Chilean, Subantarctic and Patagonian subre- gions [sensu 50]. Apart from these, all other proposals for bioregionalization for aquatic insects have been made with too large divisions [e.g., 51], making it difficult to use. Here we’ll follow the gradients of de Moor & Ivanov’ proposal [20] and when necessary subregions of Morrone and Escalante [48–50]. Distributional data and richness estimates

The distribution data basis was compiled through the primary literature (description and record of the occurrence of spe- cies), the database of Global Biodiversity Information Facility (GBIF, https://www.gbif.org), and SpeciesLink (http://www. splink.org.br/). Original data were obtained from collections at the Museu de História Natural da Bahia of the Universidade Federal da Bahia (UFBA), the Laboratório de Citotaxonomia e Insetos Aquáticos of the Instituto Nacional de Pesquisas da Amazônia (LACIA-INPA), the Museum of Comparative Zoology at Harvard University (MCZ), the University of Minne- sota Insect Collection (UMSP), and the National Museum of Natural History of the Smithsonian Institution (USNM). As the study was based on previously collected and curated material from these museums and institutional collections, no additional permits were required for the collection or transportation of biological specimens. We use gazetteers and Google Maps© to register localization without coordinates; the centroid of the less comprehen- sive location was used. After the data compilation, two-stage filtering process was performed, (i) manual selection of the data, discarding points without coordinate information, generic data of locality (e.g., only state), or with the indeterminacy of the species (e.g., identification only to the genus level), and (ii) selection in R environment, discarding points that can generate an analysis bias (e.g., points localized at the centroid of the capital areas and/or with same coordinates or in

marine areas). Species distribution map and heatmap (Kernel density) were prepared using QGIS v. 3.10.10. Distribu- tional records of 146 species, of which 17 species are H. (Cochliopsyche) and 129 species are H. (Feropsyche) (Table 1; S1 Table) and five undescribed species of H. (Cochliopsyche) (being described in another paper) were catalogued. For estimating the number of unknown species of H. (Feropsyche) in NA-NT-PA and H. (Cochliopsyche) in NT, the distri- butional records and bioregions delimited here are used, using non-parametric estimators. Estimators were calculated based on incidence data (presence-absence only), using freshwater ecoregions (Abell et al. 2008) as sampling unities, counting the occurrences in each one, with the function ‘specpool’ from the ‘vegan’ package [52] in the R environment. This function calculates two estimators of species richness: CHAO2, second-order jackknife (JACK2) (more adequate for incidence data) [53]. These non-parametric estimators are useful to estimate a potential number of unobserved species based on incidence data as those available here, and they have shown better performance than model-based or asymptotic estimators [54,55]. Environmental gradients To obtain the standards of distribution and distribution range regarding temperature, precipitation, elevation, and char- acteristics of freshwater environments, a dataset was produced consisting of distribution data, biogeoclimatic data (i.e.,

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 5 / 22 Table 1. Species of Helicopsyche and information of known semaphoronts, distribution, and collections with deposited material. Species KS Museum Distribution H. (C.) amazona Johanson, 2003 ♂ USNM, NRM BRA H. (C.) amica Johanson, 2003 ♂, ♀* MVC, USNM, MCZ, ISNB BRA, GUY, VEN H. (C.) blahniki Johanson, 2003 ♂, ♀* UMSP, CIUC, FMNH, MVC, USNM NRM BRA, COL, ECU, GUY, PER, VEN H. (C.) brazilia Johanson, 2003 ♂, ♀* MZUSP, USNM, MRM BRA H. (C.) chocoensis Johanson, 2003 ♂, ♀* USNM BRA, COL H. (C.) clara (Ulmer, 1905) ♂, ♀ MCZ, MZUSP, UMSP, USNM ARG, BRA, ECU H. (C.) holzenthali Johanson, 2003 ♂, ♀* UMSP, USMN VEN H. (C.) lobata (Flint, 1983) ♂ USNM, MCZ, MZUSP, UMSP ARG, BRA, PER H. (C.) nyurga Oláh & Oláh, 2022 ♂ OPC ECU H. (C.) napoa Johanson, 2003 ♂, ♀* USNM ECU H. (C.) ocosingua Johanson, 2003 ♂, ♀* INHS, NRM BRA, MEX H. (C.) opalescens (Flint, 1972) ♂ USNM, FNMH, MZUSP, UMSP ARG, BRA, ECU, GUY, PAR, PER, SUR, URU, VEN H. (C.) pandeirosa Johanson, 2003 ♂, ♀* MZUSP, UMSP, NRM, USNM BRA H. (C.) paraguaiensis Johanson, 2003 ♂ USNM PAR H. (C.) puyoa Johanson, 2003 ♂, ♀* USNM, UMSP BRA, ECU H. (C.) vazquezae (Flint, 1986) ♂, L, P, C USNM, INHS, UMSP BOL, COR, ECU, MEX, VEN H. (C.) xinguensis Johanson, 2003 ♂, ♀* MZUSP, USNM, UMSP BRA H. (F.) alajuela Johanson & Holzenthal, 2010 ♂ NMNH; COZEM CRI; PAN H. (F.) altercoma Botosaneanu & Flint, 1991 ♂♀ NMNH; CMNH; FSCA; ZMUA DOM

H. (F.) angeloi Holzenthal, Blahnik & Calor, 2016 ♂♀* MZUSP; UFBA; UMSP BRA H. (F.) angulata Flint, 1981 ♂♀ USNM; UMSP; NRM COL; ECU; VEN H. (F.) apicauda Flint, 1968 all USNM; NMNH DMA; GUA H. (F.) auroa Johanson & Holzenthal, 2004 ♂♀* UMSP; NMNH VEN H. (F.) bendego Dumas & Nessimian, 2019 ♂♀* DZRJ BRA H. (F.) blancasi Schmid, 1958 ♂♀* NMNH PER H. (F.) blantoni Johanson & Malm, 2006 ♂ NMNH; NRM PAN H. (F.) borealis (Hagen, 1861) all MCZ; UMSP; NMNH; TAMU; USNM CAN; CRI; GTM; HND; MEX; NIC; PAN; USA H. (F.) braziliensis (Swainson, 1840) C – BRA H. (F.) breviterga Flint, 1991 ♂♀* UMSP; COZEM; UMSP; NRM COL; PAN; VEN H. (F.) caligata Flint, 1967 ♂ NMNH CHI H. (F.) camuriensis Johanson & Holzenthal, 2004 ♂♀* UMSP VEN H. (F.) carajas Gama Neto, Ribeiro & Passos, 2019 ♂ MPEG BRA H. (F.) catoles Souza, Gomes & Calor, 2017 ♂♀* MZUSP; UFBA; UFRJ BRA H. (F.) centrocubana Botosaneanu & Flint, 1991 ♂L ZMUA CUB H. (F.) chilensis Flint, 1983 ♂♀* NMNH CHI H. (F.) chiriquensis Johanson & Malm, 2006 ♂♀* NMNH; UMSP; INBIO CRI; PAN H. (F.) cipoensis Johanson & Malm, 2006 ♂ NMNH BRA H. (F.) circulata Johanson & Holzenthal, 2004 ♂ UMSP VEN H. (F.) cochleara Johanson, 1999 ♂ NMNH ECU H. (F.) colombiensis von Siebold, 1856 C – COL; VEN H. (F.) comosa Kingsolver, 1964 ♂♀ INHS; ZMUA; NMNH; MCZ CUB H. (F.) cotopaxi Botosaneanu & Flint, 1982 ♂♀*L*P*C USNM; ZMUA ECU (Continued)

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 6 / 22 Species KS Museum Distribution H. (F.) cubana Kingsolver, 1964 all* INHS; NHMJ; ZMUA CUB; JAM H. (F.) curvipalpia Johanson & Malm, 2006 ♂♀* INHS; NRM MEX H. (F.) dampfi Ross, 1956 ♂♀*P* INHS; CNHM; NMNH; UMSP; INBIO; MEL CRI; GTM; MEX; NIC H. (F.) daome Dumas & Nessimian, 2019 ♂♀* DZRJ BRA H. (F.) diamantina Pereira & Calor, 2024 ♂ MZUSP; UFBA BRA H. (F.) dinoprata Dumas & Nessimian, 2019 ♂♀* DZRJ; MZUSP BRA H. (F.) disjuncta Johanson & Holzenthal, 2004 ♂ NMNH VEN H. (F.) dominicana Botosaneanu & Flint, 1991 ♂♀ USNM; CMNH; CNHM; FSCA; NMNH; ZMUA DOM H. (F.) dorsocurvata Johanson & Holzenthal, 2010 ♂ UMSP CRI † H. (F.) electra Johanson & Wichard, 1996 ♂ Collection Wichard DOM H. (F.) extensa Ross, 1956 ♂♀* INHS; UMSP PER; VEN H. (F.) falcigona Botosaneanu & Flint, 1991 ♂♀*L ZMUA; USNM; MCZ CUB H. (F.) fistulata Flint, 1991 ♂♀* USNM COL; VEN H. (F.) flinti Johanson, 1999 ♂ BMNH BRA H. (F.) fridae Johanson, 2003 ♂♀* NMNH; UCD PAN H. (F.) golfitoensis Johanson & Holzenthal, 2010 ♂ NMNH CRI H. (F.) granpiedrana Botosaneanu & Sykora, 1973 ♂ ZMUA CUB H. (F.) grenadensis Flint & Sykora, 1993 ♂♀* FSCA; NMNH; UMSP; NRM GRE; VEN H. (F.) guadeloupensis Malicky, 1980 all CM; ZMUA; MHNH; CMNH; MNNM; MNHM DMA; GUA; LCA; MTQ H. (F.) guara Holzenthal, Blahnik & Calor, 2016 ♂♀* MZUSP; UMSP; UFBA BRA H. (F.) guariru Vilarino & Calor, 2017 ♂ MZUSP; UMSP; UFBA BRA H. (F.) hageni Banks, 1938 all MCZ; ZMUA CUB; DOM

H. (F.) haitiensis Banks, 1938 ♂ MCZ HTI H. (F.) helicoidella (Vallot, 1855) C – BRA H. (F.) incisa Ross, 1956 ♂♀* INHS; UMSP; INBIO; NMNH; NRM CRI; MEX; NIC; PAN H. (F.) imperial Silva-Pereira, Desidério, Pereira, Hamada, 2024 ♂ INPA, UFBA BRA H. (F.) johansoni Moreno, Desidério, Pes & Hamada, 2023 ♂ INPA; DZRJ; MNRJ; UFBA BRA H. (F.) inflata Gama Neto, Ribeiro & Passos, 2019 ♂ MPEG BRA H. (F.) kalaom Botosaneanu, 1996 ♂♀* ZMUA; FSCA; NMNH; CMNH DOM H. (F.) kingstona Johanson, 2003 ♂ UCD JAM H. (F.) krenak Bonfá-Neto, Vilarino & Salles, 2023 ♂ UFVB BRA H. (F.) lambda Flint, 1983 ♂ NMNH ARG H. (F.) laneblina Johanson & Holzenthal, 2004 ♂ NMNH VEN H. (F.) lara Johanson & Holzenthal, 2004 ♂♀* UMSP; IZAM; NRM VEN H. (F.) lazzariae Holzenthal, Blahnik & Calor, 2016 ♂ MZUSP BRA H. (F.) lewalleni Denning & Blickle, 1979 ♂♀ CAS; INBIO; UMSP CRI; ELS H. (F.) limnella Ross, 1937 ♂ INHS USA H. (F.) linabena Johanson & Holzenthal, 2004 ♂♀* NMNH VEN H. (F.) linguata Johanson & Malm, 2006 ♂ NMNH PAN H. (F.) lutea (Hagen, 1961) ♀ MCZ DOM H. (F.) luziae 17 ♂♀* DZRJ BRA Table 1. (Continued) (Continued)

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 7 / 22 Species KS Museum Distribution H. (F.) maculisternum Botosaneanu, 1993 ♂♀* ZMUA VEN; TRI H. (F.) manaos Moreno, Desidério, Pes & Hamada, 2023 ♂ NMNH; ZMUA; CMNH; FSCA DOM H. (F.) melanochaeta Flint & Sykora, 2004 ♂♀* INPA; DZRJ; MNRJ; UFBA BRA H. (F.) merida Botosaneanu & Flint, 1982 ♂♀*L*P*C NMNH; ZMUA; UMSP VEN H. (F.) mexicana Banks, 1901 ♂♀ MCZ; INHS; NMNH; USNM; CAS; OSU; UCR CRI; MEX; USA H. (F.) minima von Siebold, 1856 all USNM; ZMUA NIC; PRI H. (F.) minuscula Martynov, 1912 ♀ PAN PER H. (F.) molesta Botosaneanu, 1998 ♂ ZMUA JAM H. (F.) mateusi Pereira & Calor, 2024 ♂ MZUSP; UFBA BRA H. (F.) miltonsantosi Pereira & Calor, 2024 ♂ MZUSP; UFBA BRA H. (F.) monda Flint, 1983 ♂ USNM; DZRJ; NMNH ARG; BRA; PRY; VEN H. (F.) montana Felber, 1912 LPC NMB MEX H. (F.) muelleri Banks, 1920 ♂♀*LPC MCZ; IRSNB ARG; BRA; PER H. (F.) neblinensis Johanson & Holzenthal, 2004 ♂♀* NMNH; IZAM; NRM VEN H. (F.) nigrisensilla Botosaneanu & Flint. 1991 ♂♀ USNM; ZMUA DOM H. (F.) obscura Rueda Martín & Isa Miranda, 2015 ♂LPC IBN ARG H. (F.) occidentale Botosaneanu & Flint, 1991 ♂L USNM; ZMUA CUB H. (F.) ochthephila Flint, 1968 all NMNH JAM H. (F.) paprockii Johanson & Malm, 2006 ♂ NMNH BRA H. (F.) parahageni Flint & Sykora, 2004 ♂♀ NMNH; CMNH; FSCA DOM H. (F.) paralimnella Hamilton, 1989 ♂ CUEC USA H. (F.) paucispina Botosaneanu & Flint, 1991 ♂ ZMUA CUB H. (F.) perija Johanson & Holzenthal, 2004 ♂ UMSP VEN H. (F.) peruana Banks, 1920 ♂ MCZ PER

H. (F.) paulofreirei Pereira & Calor, 2024 ♂ MZUSP; UFBA BRA H. (F.) petri Dumas & Nessimian, 2019 ♂ DZRJ; MZUSP BRA H. (F.) pietia Denning, 1964 ♂♀ CAS; INHS; NRM MEX; USA H. (F.) piroa Ross, 1944 ♂♀* INHS; TAMU; USNM CRI; MEX; NIC; USA H. (F.) planata Ross, 1956 ♂ INHS; CNIN; CUEC NIC; MEX H. (F.) planorboides Machado, 1957 all DZRJ BRA H. (F.) poliochaeta Flint & Sykora, 2004 ♂♀ NMNH; FSCA DOM H. (F.) propinqua Botosaneanu & Flint, 1991 ♂ NMNH PRI H. (F.) quadrosa Ross, 1956 ♂ INHS MEX H. (F.) ralphi Cavalcante-Silva, Pereira & Calor, 2022 all MZUSP; INPA; UFBA; UFRJ BRA H. (F.) ramosi Flint, 1964 all NMNH; ZMUA PRI H. (F.) rentzi Denning & Blickle, 1979 ♂♀ CAS; INBIO; UMSP; USNM CRI H. (F.) sanblasensis Johanson & Malm, 2006 ♂ NMNH PAN H. (F.) scalaris Hagen, 1864 C – VEN † H. (F.) electra Johanson & Wichard, 1996 ♂ Collection Wichard DOM H. (F.) selanderi Ross, 1956 ♂♀* INHS; NMNH; UMSP CRI; MEX; VEN H. (F.) septifera Flint & Sykora, 2004 ♂♀ NMNH; CMNH; ZMUA DOM H. (F.) shaamunensu Dumas & Nessimian, 2019 ♂ DZRJ BRA H. (F.) sigillata Botosaneanu & Flint, 1991 ♂♀ NMNH; ZMUA CUB Table 1. (Continued) (Continued)

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 8 / 22 Species KS Museum Distribution H. (F.) singulare Botosaneanu & Flint, 1991 ♂ NMNH; USNM; ZMUA PRI H. (F.) sinuata Denning & Blickle, 1979 ♂ UCD; NMNH MEX; USA H. (F.) succincta Johanson & Holzenthal, 2004 ♂ NMNH; UFBA BRA; VEN H. (F.) sucrensis Johanson & Holzenthal, 2004 ♂ UMSP VEN H. (F.) tachira Johanson & Holzenthal, 2004 ♂ UMSP VEN H. (F.) tapadas Denning, 1966 ♂ CAS; UFBA BRA; VEN H. (F.) temora Denning & Blickle, 1979 ♂ UCD MEX H. (F.) thelidomus Hagen, 1864 C – VEN H. (F.) timbira Silva, Santos & Nessimian, 2014 ♂♀* DZRJ; MNRJ; INPA BRA H. (F.) truncata Ross, 1956 ♂ INHS; UMSP; USNM CRI; MEX; PAN H. (F.) turbida Navás, 1923 all MZBS; NMNH; IBN; USNM ARG H. (F.) tuxtlensis Bueno-Soria, 1983 ♂♀* IBUNAM; UCD; USNM GTM; MEX; PAN H. (F.) umbonata Hagen, 1864 all USNM; MCZ; AMNH; USNM JAM H. (F.) valligera Flint, 1983 ♂♀* NMNH; USNM ARG; BRA H. (F.) venezuelensis Johanson & Holzenthal, 2004 ♂ UMSP VEN H. (F.) vergelana Ross, 1956 all INHS; NMNH; NRM; UCD; UFPE; UMSP; USNM BLZ; BRA; COL; CRI; ECU; GTM; GUY; MEX; NIC; PAN; PER; SUR; TRI; VEM H. (F.) villegasi Denning & Blickle, 1979 ♂ UCD MEX † H. (F.) voigti Johanson & Wichard, 1996 ♂ Collection Wichard DOM H. (F.) woldai Johanson, 2003 ♂ NMNH; UCD PAN H. (F.) woytkowskii Ross, 1956 ♂♀* INHS; NRM; UMSP PAN; PER; VEN KS = Known semaphoronts; ♂ = Male; ♀ = Female; L = Larvae; P = Pupa; Collections that house type material in bold; *semaphoronts present in material

examined of the publications, but not formally described. https://doi.org/10.1371/journal.pone.0327580.t001 Table 1. (Continued) temperature, precipitation, elevation) obtained through WorldClim version 2.1 (https://www.worldclim.org/data/world- clim21.html), and data of freshwater ecosystems (i.e., Sthaler order, flow velocity, discharge, water body size) available in HydroRIVERS and HydroLAKES version 1.0 (https://www.hydrosheds.org). Based on this, distribution value plots and boxplots were generated using the ‘ggplot’ function of the ggplot2 package, a correlation analysis between the variables was performed with the Spearman method using the ‘correlate’ function of the ‘corrr’ package (https://cran.r-project.org/ package=corrr) and PERMANOVA analysis with Jaccard similarity index was performed to assess the similarity between subgenus both performed in R environment. Distribution modelling Environmental data were obtained from monthly climate data for minimum, mean, and maximum temperature, precipita- tion, solar radiation, wind speed, water vapor pressure, and for total precipitation, 19 “bioclimatic” variables, and elevation on a scale of 5 arc minutes, all environmental variables obtained in WorldClim version 2.1 (https://www.worldclim.org/ data/worldclim21.html). The resolutions of these environmental variables were kept, avoiding the loss of information or the impossibility of modelling. After obtaining the data, a correlation analysis between the variables was performed with the

Spearman method using the ‘correlate’ function of the ‘corrr’ package (https://cran.r-project.org/package=corrr) in the R environment. This correlation analysis permitted to selection of uncorrelated variables and consequently to avoid over- weighing in the models. Variables with correlation values greater than 80% were considered correlated.

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 9 / 22 Four correlative modelling algorithms were used: Bioclim [56], Domain [57], Generalized Linear Model (GLM) [58], and the Vector Support Machine (SVM) [59]. As absence data is not predicted or lacks accuracy given the bias/scarce sam- pling scenario, presence/pseudo-absence models will be used. We randomly generated pseudo-absence points (1:1 ratio for the occurrence points) [60], through the ‘randomPoints’ function of the ‘dismo’ package (https://cran.r-project.org/pack- age=dismo) in the R environment. Data partitioning was randomly performed in 70% for training and 30% for testing the models. The repeatability of the models (200 times) was used to increase the robustness of the result. After the models were evaluated using the Area Under the Curve (AUC) [61] method, which only models with values of AUC > 80% were used for the construction of suitability maps. Finally, we used the default limits of presence and absence for the construc- tion of the suitability maps. Results Biodiversity knowledge shortfalls In the New World, the Helicopsyche species richness remains largely underestimated, with estimates suggesting up to 571 species (error = 142), indicating that approximately 75% of the species remain undescribed (Table 2). The subgenus H. (Cochliopsyche) comprises 17 valid species, all distributed in the Neotropics, particularly in the Brazilian subregion (Fig 1B). Estimates project 36 species (error = 16), with 53% remaining undescribed (Table 2). The first species was described

by Ulmer in 1905, followed by significant contributions from Flint [62–64]. Johanson’s revision added 12 species, and after a 20-year gap, Oláh & Oláh [65] described a new species from Ecuador (Fig 2C). The subgenus H. (Feropsyche) includes 129 valid species, widely distributed across tropical regions of the Nearctic, Neotropical and Patagonian regions, with the highest richness in the Brazilian subregion (125 species) (Fig 1G). Estimates suggest 553 species (error = 152), with 77% remaining undescribed (Table 2). In the past 20 years, about 40% of H. (Feropsyche) species were described, averaging five species annually, compared to two species per year from 1840 to 1999 (Fig 2D). Approximately 79% of Helicopsyche species have restricted distributional records (< 10 records), with around 70% occurring only in type localities or adjacent areas, such as the same stream (Fig 2A and B; S1 Table). The species with the widest distribution are H. (Cochliopsyche) opalescens Flint, 1972, with 41 records across 147.3 km², and H. (Feropsy- che) borealis (Hagen, 1861), with 48 records across 211.03 km² (Fig 2A and B). In H. (Cochliopsyche), 58% of species have restricted records, with 47% limited to type localities or adjacent areas (Fig 2A; S1 Table). In contrast, H. (Feropsy- che) shows a higher percentage of species with restricted records, reaching 88%, and 72% of these species are confined to type localities or adjacent areas (Fig 2B; S1 Table). Immature stages are documented for 20 valid species (15%), and female adults are described for 30 valid species

(24%). In H. (Cochliopsyche), adult males of all species are described. However, immature stages are known for only one Table 2. Summary of the estimation of species richness in the new world based on incidence data with the CHAO2, second order jackknife (JACK2) estimators. Chao2 Jacknife2 Average Estimation of unknown species Helicopsyche (Cochliopsyche) Species estimates 36 34 35 18 Error 16 4 10 Helicopsyche (Feropsyche) Species estimates 553 310 431 302 Error 152 33 92 New World Helicopsyche Species estimates 571 343 457 311 Error 142 35 88 https://doi.org/10.1371/journal.pone.0327580.t002

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 10 / 22 species, H. (Cochliopsyche) vazquezae (Flint, 1986), and females are documented for two species, H. (Cochliopsyche) clara (Ulmer, 1905) and H. (Cochliopsyche) vazquezae (Table 1). In H. (Feropsyche), males of 121 species and females of 28 species are described, including two species known solely from females: H. (Feropsyche) lutea (Hagen, 1861) and H. (Feropsyche) minuscula Martynov, 1912. Immature stages have been documented for 19 species of H. (Feropsyche). Fig 1. Distribution and environmental suitability map for the New World Helicopsychidae species. A. Environmental suitability map (weighted average) of Helicopsyche (Cochliopsyche); B. Distribution records of Helicopsyche (Cochliopsyche) species; C. Map of high environmental suitability (>0.7) of Helicopsyche (Cochliopsyche); D. Environmental suitability map (weighted average) of Helicopsyche (Feropsyche); E. Distribution records of Helicopsyche (Feropsyche) species; F. Map of high environmental suitability (>0.7) of Helicopsyche (Feropsyche); G. Distribution records of New World Helicopsychidae species; H. Map of high environmental suitability (>0.7) of New World Helicopsychidae. Map created in QGIS v3.40.7. Background: ESRI land cover (https://livingatlas.arcgis.com/landcover/) accessed via the QuickMapServices plugin, country boundaries: geoBoundaries (https://www. geoboundaries.org/), both under CC BY 4.0 license. https://doi.org/10.1371/journal.pone.0327580.g001

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 11 / 22

has 74% of its records in areas above 1000 mm³/year (range 959–3798 mm³/year) (Fig 4B), with significant differences (F(1)=58.52, p-value = 0.001). Temperature gradients range from −17°C to 40°C. The H. (Feropsyche) species is the only subgenus found in areas with negative temperatures, with a range from −17°C to 40°C (Fig 4C). In contrast, H. (Cochlio- psyche) species are found between 5°C and 35°C (Fig 4D), with significant differences (F(1)=55.10, p-value = 0.001). Distribution patterns of New World The northernmost species of H. (Feropsyche) is H. (Feropsyche) borealis, recorded in Canada at approximately 47°N, while the southernmost is H. (Feropsyche) caligata, found in Chile at around 39°S. In H. (Cochliopsyche), the northern- most species is H. (Cochliopsyche) vazquezae from Mexico (17°N), and the southernmost is H. (Cochliopsyche) opal- escens from Uruguay (32°S). The highest species richness in both subgenera is concentrated between latitudes 0° and 24°S (Fig 1). In the New World, biodiversity hotspots H. (Feropsyche) are concentrated in three main regions: (i) northwestern Colombia extending through Costa Rica, Nicaragua, and Panama, with 231 records representing 25 species; (ii) the Greater Antilles, with 165 records of 22 species; and (iii) eastern Brazil, with 93 records of 13 species (Fig 2A). For H. (Cochliopsyche), three prominent hotspots include: (i) northern Peru, much of Colombia, and Ecuador, with some exten- sion into the Amazon basin; (ii) the Central Amazon in Brazil; and (iii) eastern Brazil, encompassing the Central Atlantic

Forest Ecological Corridor and adjacent areas, and high-altitude areas of the Caatinga domain (Fig 2B). Species of Helicopsyche occur across 90 freshwater ecoregions [sensu 66]. The Northeastern Mata Atlântica Fresh- water Ecoregion harbors the highest species richness (23 spp.), followed by the South America Caribbean Freshwater Ecoregion (15 spp.), and both the Amazonas Lowlands and Paraíba do Sul Freshwater Ecoregions (14 species each). Helicopsyche (Cochliopsyche) is predominantly recorded from the Amazonas Lowlands (10 spp.) and Northeastern Mata Atlântica (8 spp.) Freshwater Ecoregions, whereas H. (Feropsyche) is more frequent in the South America Caribbean (15 spp.), San Juan (Nicaragua and Costa Rica), and Paraíba do Sul (12 species each) Freshwater Ecoregions (see S2 Table for details). Fig 2. Distribution range and species new species history of New World Helicopsychidae. A. Number of records and distribution range of Heli- copsyche (Cochliopsyche) species. B. Number of records and distribution range of Helicopsyche (Feropsyche) species. C. History of new and number of Helicopsyche (Cochliopsyche) species over time. D. History of new and number of Helicopsyche (Feropsyche) species over time. E. Heatmap of distribution for Helicopsyche (Cochliopsyche). F. Heatmap of distribution for Helicopsyche (Feropsyche). Map created in QGIS v3.40.7. Background: ESRI land cover (https://livingatlas.arcgis.com/landcover/) accessed via the QuickMapServices plugin, country boundaries: geoBoundaries (https://www. geoboundaries.org/), both under CC BY 4.0 license.

https://doi.org/10.1371/journal.pone.0327580.g002

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 13 / 22

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 14 / 22 Distribution modelling After conducting correlation testing, nine raster variables were found to be uncorrelated, belonging to five groups (biocli- matic, elevation, precipitation, solar radiation, and wind speed), as listed in Table 3 (for more details, see S3 Table and S1 Fig). Among the four algorithms tested, all yielded AUC values higher than the cut-off value (S2 Fig). These values were subsequently used to generate environmental suitability maps for the subgenera (Fig 1). Distribution modelling indicates high environmental suitability (values > 0.7) for H. (Cochliopsyche) species across broad areas of the Neotropical region, particularly throughout the Brazilian and Chacoan subregions. Areas of notable suitability include: (i) the Amazon Basin, (ii) the Atlantic Forest dominion, and (iii) high-altitude regions in the eastern por- tions of the Caatinga and Cerrado dominions (Fig 1A and C). In contrast, areas of low suitability are primarily found in the (i) Mexican and South American transition zones; (ii) the Mesoamerican dominion, and (iii) the Antillean subregion. Within the Brazilian subregion, low suitability is also observed in the (iv) Pará, Xingu-Tapajos, Cerrado, Chacoan, and Pampean provinces, as well as across the entire (v) Patagonian region (Fig 1A and C). For H. (Feropsyche) species, the model predicts high environmental suitability (> 0.7) primarily in the Nearctic region,

especially: (i) in South Florida (Florida Keys), where no species records currently exist. In the Mexican transition zone, (ii) southern portions of the Chihuahuan Desert, Sierra Madre Occidental and Oriental, and the Trans-Mexican Volcanic Belt also show high suitability. In the Neotropical region, four major areas exhibit high suitability: (iii) the entire Antillean subregion; in the Brazilian subregion (iv) most part of the Mesoamerican dominion, excluding the coastal areas of the Veracruzan province and nearly all of the Yucatán Peninsula; (v) in the Pacific dominion, the Guatuso–Talamanca, Puntar- enas–Chiriquí, and Chocó–Darién provinces, as well as high-altitude (Andean Cordillera) and coastal regions of Colombia and Venezuela—except much of the Guajira and Magdalena provinces; (vi) in the Boreal Brazilian dominion, the western portions of the Guiana Lowlands and Yungas provinces; and Chacoan subregion (vii) the entire Paraná domain and areas east of Caatinga Cerrado and Chacoan provinces where they are bordered by the Atlantic Forest and Paraná provinces (Fig 1D and F). Conversely, areas of low environmental suitability for H. (Feropsyche) species are primarily located in: (i) most of the South American transition zone; (ii) nearly all the Boreal Brazilian, South Brazilian, and Southeastern Amazo- nian dominions; and (iii) most of the Chacoan dominion, except for eastern portions of Caatinga, Cerrado and Chacoan provinces (Fig 1D and F). Discussion Biodiversity knowledge shortfalls and challenges

Our results demonstrate the clear increment of species descriptions and cataloguing of distributional records, mainly in under-explored areas of Neotropical region. This increase can be related with the development of trichopterology in the Neotropical, Nearctic and Patagonian regions [as indicated in 33,67], cooperation among researchers of NT with other researchers’ groups [e.g., 24,36], and the establishment of research groups in different countries of the Neotropical and Patagonian regions [e.g., 23,38,68]. After Johanson [30,35,41,45,69], who established the modern panorama of Helico- psychidae taxonomy in the Neotropical, Nearctic and Patagonian regions, several contributions have been made by local researchers [e.g., 32,37,40,62,70], but no one with a broad perspective. However, our results indicate more dramatic scenarios for Helicopsyche, with up to 75% of the fauna still unknown and numerous unexplored areas, when compared with previous estimates for Trichoptera in the NT [23,24], both with around 50% unknown species] (Table 2). These results are consistent when looking at the number of known species and the vast Fig 3. Boxplot of biogeoclimatic variables of New World Helicopsyche species. A. Elevation range of Helicopsyche (Cochliopsyche) species; B. Elevation range of Helicopsyche (Feropsyche) species; C. Freshwater Stahler order range of Helicopsyche (Cochliopsyche) species; D. Freshwater Stahler order range of Helicopsyche (Feropsyche) species. https://doi.org/10.1371/journal.pone.0327580.g003

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 15 / 22

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 16 / 22 areas with high suitability and little or no research effort, so efforts should be made to describe new species. In addition to unknown species, some valid species exhibit weak circumscription or inconsistencies [see 32,40]. In H. (Cochliopsyche), there is little difference in genital characteristics (the primary features for species distinction), thus, detailed description of genitalia, standardization, and better circumscription of species are relevant to avoid misidentifications. The Wallacean shortfall is evident in the limited data available on species distribution [8]. Distributional information is often restricted to type localities or adjacent areas (Fig 2A and B), a trend that is also reflected in our results. This high- lights the urgent need to increase faunistic surveys in unexplored areas with high environmental suitability (Fig 1C,F, and H) and to analyze the material already housed in collections. Key regions with significant research effort and species rich- ness include: (i) Antillean subregion in Hispaniola, Puerto Rico – Virgin Islands, Windward & Leeward Islands, and Cuba Cayman Islands (eastern and western extremes) Freshwater Ecoregions; (ii) Mesoamerican dominion in San Juan, Estero Real – Tempisque, Chiriqui, Isthmus Caribbean, Chagres, South America Caribbean Drainages – Trinidad and south portion of Magdalena – Sinu Freshwater Ecoregions, (iii) Brazilian dominion only near Manaus city, Amazonas, and (iv)

Parana dominion in Paraíba do Sul, Northeastern Mata Atlântica, São Francisco, and Fluminense Freshwater Ecoregions (Fig 1G and H). In contrast, areas such as (i) Boreal and South Brazilian dominions; and (ii) areas east of Caatinga Cerrado and Chacoan provinces where they are bordered by the Atlantic Forest and Paraná provinces remain largely underex- plored despite showing high environmental suitability, indicating strong potential for species distribution. Prioritizing these regions for future faunistic surveys targeting Helicopsychidae is essential to fill existing knowledge shortfalls. These efforts will likely lead to the discovery of previously undescribed taxa and a more comprehensive understanding of species distributions [8]. In doing so, they will help address both Linnean and Wallacean shortfalls, while also con- tributing to resolving Haeckelian and Hutchinsonian shortfalls if semaphoronts and abiotic conditions are thoroughly documented. Given the Wallacean and Linnean shortfalls in caddisfly research, our understanding of species diversity and taxo- nomic groups remains incomplete, particularly within Trichoptera [36], where a male-biased focus persists—a gap that aligns with the Haeckelian shortfall [more details, 9]. Biodiversity biases often stem from research concentrated in easily accessible areas with prior survey efforts [3]; however, in this case, the sexual bias is significantly amplified. Therefore, acknowledge that all subsequent discussions are inherently influenced by these knowledge shortfall, reflecting the broader

discrepancies in our understanding of New World Helicopsychidae, a scenario that is similarly prevalent across most insect orders. Fig 4. Boxplot of biogeoclimatic variables of New World Helicopsyche species. A. Annual mean precipitation range of Helicopsyche (Cochliopsy- che) species; B. Annual mean precipitation range of Helicopsyche (Feropsyche) species; C. Annual mean temperature range of Helicopsyche (Cochlio- psyche) species; D. Annual mean temperature range of Helicopsyche (Feropsyche) species. https://doi.org/10.1371/journal.pone.0327580.g004 Table 3. Environmental variables used for species modelling, using 80% correlation cut-off. Variables group Code Variables Bioclimatic Bio_7 Temperature Annual Range (Annual Precipitation- Min Temperature of Coldest Month) Bio_2 Annual Precipitation Elevation Elev Elevation Precipitation Prec_01 and 08 Precipitation (mm) Solar radiation Srad_03, 04 and 07 Solar radiation (kj m-2 day-1) Wind speed Wind_01 Wind speed (m s-1) https://doi.org/10.1371/journal.pone.0327580.t003

PLOS One | https://doi.org/10.1371/journal.pone.0327580 July 28, 2025 17 / 22 The H. (Feropsyche) has 36 valid species, and H. (Cochliopsyche) present ten valid species with adult females depos- ited (in material examined), but not formally described in published studies (Table 1). Both subgenera have specimens of immature stages deposited in museums and collections that have not yet been associated with adult males (Table 1). These data demonstrate the enormous potential of the material to be described and fulfil the biodiversity shortfalls. The description of these semaphoronts can contribute to a better circumscription of the species, the correction of possible taxonomic errors, and the formation of a complementary database for studies related to the understanding of relationships and biogeographic history [9]. To our knowledge, this is one of first inference describing environmental gradients for Trichoptera species [preceded by the work of [25,26], which investigating the influence of elevation on Trichoptera]. Here are addresses Hutchinsonian deficits by empirically defining species’ abiotic limits and identifying areas with a higher probability of distribution in regions of high suitability, focusing on the niches of H. (Feropsyche) and H. (Cochliopsyche). This also provides evidence of niche differentiation between the two living lineages of Helicopsychidae in the New World. Additionally, these data help identify species more vulnerable to extinction due to their narrow distribution gradients in biogeoclimatic variables. However, we