Abstract

The Northeastern Mata Atlântica Freshwater ecoregion (NMAF) is part of the 25 worlds biodiversity hotspots. It comprises the Central Atlantic Forest Ecological Corridor and Chapada Diamantina Complex (in part), including high rates of endemism in coastal freshwater ecosystems. However, estimates indicate a high population decline in Freshwater ecosystems. Trichoptera are the most affected insect order, with average extinction rates of ~9% and many unknown species (e.g., estimates are around 50% in Brazil and Ecuador). This crisis can be aggravated by gaps in the knowledge of species (Linnean shortfall) and their distribution (Wallacean shortfall), caused mainly by a lack of investment in extensive fauna inventories and human resources related to systematics. Thus, to face these shortfalls in NMAF, we describe four new species of. H. (Feropsyche) and provide new distribution records. In addition, we perform niche modeling based on the species distributions of the group to identify areas with high environmental suitability to direct biodiversity research efforts on NMAF, a highly endemic and underexplored ecoregion. We increased the number of known species of NMAF from seven to 16 species. The niche modeling pointed to two areas as priorities to guide the strategies to reduce shortfalls in the NMAF.

Key words
Atlantic Forest; biodiversity shortfalls; potential distribution; Helicopsychidae; snail-case caddisflies

INTRODUCTION

Freshwater ecosystems cover around 1% of Earth’s land surface but comprise ca. 10% of all known species (Strayer & Dudgeon 2010STRAYER DL & DUDGEON D. 2010. Freshwater biodiversity conservation: recent progress and future challenges. J North Amer Benthol Soc 29: 344-358. https://doi.org/10.1899/08-171.1.
https://doi.org/10.1899/08-171.1... ), with a high rate of endemism (Watson et al. 2018WATSON JEM ET AL. 2018. The exceptional value of intact forest ecosystems. Nat Ecol Evol 2: 599-610. https://doi.org/10.1038/s41559-018-0490-x.
https://doi.org/10.1038/s41559-018-0490-... ). These species have experienced significant population declines compared to terrestrial and marine ecosystems (MEA 2005MEA - MILLENIUM ECOSYSTEM ASSESSMENT. 2005. Ecosystems and human well-being, v. 5, p. 563. United States of America: Island press.). Due to these characteristics, freshwater ecosystems have the most acute biodiversity crisis among ecosystems (Tickner et al. 2020TICKNER D ET AL. 2020. Bending the Curve of Global Freshwater Biodiversity Loss: An Emergency Recovery Plan. BioScience 30: 330-342. https://doi.org/10.1093/biosci/biaa002.
https://doi.org/10.1093/biosci/biaa002... ). In addition, biodiversity knowledge shortfalls on freshwater ecosystems are pronounced as the result of several factors such as neglected large areas, few comprehensive inventories, and lack of specialized human resources and/or investment in biodiversity research, especially in developing countries (Kier et al. 2005KIER G ET AL. 2005. Global patterns of plant diversity and floristic knowledge. J Biogeogr 32: 1-10. https://doi.org/10.1111/j.1365-2699.2005.01232.x.
https://doi.org/10.1111/j.1365-2699.2005... , Collen et al. 2008COLLEN B, RAM M, ZAMIN T & MCRAE L. 2008. The tropical biodiversity data gap: addressing disparity in global monitoring. Tropical Cons Sci 1: 35-88. https://doi.org/10.1133%2F194008290800100202.
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https://doi.org/10.1111/ddi.12489... ).

Tropical freshwater ecosystems house rich biodiversity, with faunal components more susceptible to impacts related to the degradation of the forests, pollution of freshwater environments, and climate change (Tewksbury et al. 2008TEWKSBURY JJ, HUEY RB & DEUTSCH CA. 2008. Ecology: Putting the heat on tropical animals. Science 320: 1296-1293. http://doi.org/10.1126/science.1159328.
https://doi.org/10.1126/science.1159328... , Senior et al. 2019SENIOR RA, HILL JA & EDWARDS DP. 2019. Global loss of climate connectivity. Nat Climate Change 9: 623-626. https://doi.org/10.1038/s41558-019-0529-2.
https://doi.org/10.1038/s41558-019-0529-... ). Recent estimates indicate that the population decline of aquatic insect species is twice that of vertebrates, and local extinction rates are eight times greater for insects than for vertebrates (Sánchez-Bayo & Wyckhuys 2019SÁNCHEZ-BAYO F & WYCKHUYS KA. 2019. Worldwide decline of the entomofauna: A review of its drivers. Biol Conservation 232: 8-23. https://doi.org/10.1016/j.biocon.2019.01.020.
https://doi.org/10.1016/j.biocon.2019.01... ). Currently, 33% of aquatic insect species are threatened with extinction, and each year about 1% of all species have been added to the list, resulting in an average extinction rate (species not observed in >50 years) of 9% for some groups, such as Trichoptera (Sánchez-Bayo & Wyckhuys 2019SÁNCHEZ-BAYO F & WYCKHUYS KA. 2019. Worldwide decline of the entomofauna: A review of its drivers. Biol Conservation 232: 8-23. https://doi.org/10.1016/j.biocon.2019.01.020.
https://doi.org/10.1016/j.biocon.2019.01... ). Additionally, the uncertainty about the extinction of insect species is standard because rare or highly threatened species are intrinsically difficult to detect (Ladle et al. 2011LADLE R, JEPSON P, MALHADO A, JENNINGS S & BARUA M. 2011. Perspective: The causes and biogeographical significance of species’ rediscovery. Front Biogeogr 3: 111-118.), which can be exacerbated by biodiversity shortfalls (Mulieri et al. 2022MULIERI PR, MIGALE S, PATITUCCI LD, GONZÁLES GR & MONTEMAYOR SI. 2022. Improving geographic distribution data for a putatively extinct species, a test case with a disappeared fly. An Acad Bras Cienc 94: e20201439. https://doi.org/10.1590/0001-3365202220201439.
https://doi.org/10.1590/0001-33652022202... ).

Some solutions are presented for better use of research funding and biodiversity shortfalls, such as using tools like niche modeling to identify areas of high environmental suitability (Guisan & Zimmermann 2000GUISAN A & ZIMMERMANN NE. 2000. Predictive habitat distribution models in ecology. Ecol Model 135: 143-186. https://doi.org/10.1016/S0304-380000.00354-9.
https://doi.org/10.1016/S0304-380000.003... ). Identifying these areas with a high probability of potential distribution can analyze the material deposited in museums and collections [e.g., around 26 million specimens were deposited in Brazilian scientific collections (Joly et al. 2011JOLY CA ET AL. 2011. Diagnóstico da pesquisa em biodiversidade no Brasil. Revista USP 89: 114-133.)]. Allowing the use of already collected material (Joly et al. 2011JOLY CA ET AL. 2011. Diagnóstico da pesquisa em biodiversidade no Brasil. Revista USP 89: 114-133.) and optimizing resources for focal collections in freshwater areas with high environmental suitability and without or scarce distributional records. These areas with low Trichoptera distribution records are mainly concentrated in the northeastern Atlantic Forest and dry diagonal (Santos et al. 2020SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2020. Taxonomic Catalog of the Brazilian Fauna: order Trichoptera Insecta, diversity and distribution. Zoologia (Curitiba) 33: e46392. https://doi.org/10.3897/zoologia.37.e46392.
https://doi.org/10.3897/zoologia.37.e463... ).

Trichoptera comprises the most-rich order of aquatic insects (ca. 16.800 valid species) (Morse et al. 2023) and constitutes an essential component of freshwater ecosystems, contributing with diverse ecosystem services, including its contribution to nutrients cycling, decomposition processes, trophic network, with ecosystem engineering and biological monitoring of water quality (Morse 2013MORSE JC. 2013. Biodiversity of aquatic insects. Insect Biodiver Sci Soc 1: 205-223. https://doi.org/10.1002/9381118945568.
https://doi.org/10.1002/9381118945568... , Morse et al. 2019MORSE JC, FRANDSEN PB, GRAF W & THOMAS JA. 2019. Diversity and Ecosystem Services of Trichoptera. Insects 10: 100-125. https://doi.org/10.3390/insects10050125.
https://doi.org/10.3390/insects10050125... ). Besides its relevance, the knowledge of Brazilian species is biased, with all kinds of biodiversity deficits, specially Linnean and Wallacean shortfalls (sensu Hortal et al. 2015HORTAL J, BELLO F, DINIZ-FILHO JAL, LEWINSOHN TM, LOBO JM & LADLE RJ. 2015. Seven Shortfalls that Beset Large-Scale Knowledge of Biodiversity. Ann Rev Ecol Evol Syst 46: 523-549. https://doi.org/10.1146/annurev-ecolsys-112414-054400.
https://doi.org/10.1146/annurev-ecolsys-... ). Santos et al. (2020)SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2020. Taxonomic Catalog of the Brazilian Fauna: order Trichoptera Insecta, diversity and distribution. Zoologia (Curitiba) 33: e46392. https://doi.org/10.3897/zoologia.37.e46392.
https://doi.org/10.3897/zoologia.37.e463... recently indicated that only ca. 50% of the Brazilian caddisfly species are known. Some ecoregions, especially those located in the Northeastern of Brazil, such as the Northeast Mata Atlântica Freshwater ecoregion (NMAF), present more pronounced knowledge deficits about other ecoregions with high richness (e.g., Amazon, Parana, Southeastern Atlantic Forest ecoregions). However, with substantial advances in species knowledge in the NMAF in recent years, going from about ten species (Paprocki et al. 2004PAPROCKI H, HOLZENTHAL RW & BLAHNIK RJ. 2004. Checklist of the Trichoptera (Insecta) of Brazil I. Biota Neotrop 4: 1-22. https://doi.org/10.1590/S1676-06032004000100008.
https://doi.org/10.1590/S1676-0603200400... ) to the fourth ecoregion with the most species (137 valid species) and third with a rate of endemic species (38.7% of record species are endemic) (Santos et al. 2020SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2020. Taxonomic Catalog of the Brazilian Fauna: order Trichoptera Insecta, diversity and distribution. Zoologia (Curitiba) 33: e46392. https://doi.org/10.3897/zoologia.37.e46392.
https://doi.org/10.3897/zoologia.37.e463... ), but with still a lot to be explored.

Among the Trichoptera occurring in the NMAF, the cosmopolitan genus Helicopsyche von Siebold, 1856 is virtually found in the various freshwater ecosystems (Johanson 2002JOHANSON KA. 2002. Systematic revision of American Helicopsyche of subgenus Feropsyche (Trichoptera: Helicopsychidae). Insects Syst Evol 60: 1-151.). There are two subgenera, H. (Cochliopsyche) Müller, 1885 with one species, and H. (Feropsyche) Johanson, 1998 with seven species (Santos et al. 2023SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2023. Trichoptera in Catálogo Taxonômico da Fauna do Brasil. PNUD. http://fauna.jbrj.gov.br/fauna/faunadobrasil/238. Accessed 28 January 2023.
http://fauna.jbrj.gov.br/fauna/faunadobr... ). The two subgenera can be differentiated by the tibial spur formula [1, 2, 2 in H. (Cochliopsyche) versus 2, 4, 4 in H. (Feropsyche)] and relative antennae/body length [>1.2 in H. (Cochliopsyche) versus ≤1.2 in H. (Feropsyche)] (Johanson 2002JOHANSON KA. 2002. Systematic revision of American Helicopsyche of subgenus Feropsyche (Trichoptera: Helicopsychidae). Insects Syst Evol 60: 1-151.). Of the seven species of Helicopsyche (Feropsyche) recoding in NMAF (Tab. 1), six of them were described or recorded in the last two decades (Johanson & Holzenthal 2004JOHANSON KA & HOLZENTHAL RW. 2004. Thirteen new species and new distribution records of Helicopsyche (Feropsyche) Johanson from Venezuela (Trichoptera: Helicopsychidae). Zootaxa 711: 1-40. https://doi.org/10.11646/zootaxa.711.1.1.
https://doi.org/10.11646/zootaxa.711.1.1... , Johanson & Malm 2006JOHANSON KA & MALM T. 2006. Seven new Helicopsyche (Feropsyche) Johanson, 2002 from the Neotropical region and Nearctic Mexico (Insecta: Trichoptera: Helicopsychidae). Zootaxa 1208: 1-24. https://doi.org/10.11646/zootaxa.1208.1.1.
https://doi.org/10.11646/zootaxa.1208.1.... , Holzenthal et al. 2016HOLZENTHAL RW, BLAHNIK RJ & CALOR AR. 2016. Three new species of Helicopsyche von Siebold (Trichoptera: Helicopsychidae). from Brazil. Zootaxa 4038: 344-353. http://doi.org/10.11646/zootaxa.4038.1.29.
https://doi.org/10.11646/zootaxa.4038.1.... , Souza et al. 2017SOUZA RL, GOMES V & CALOR AR. 2017. A new species of snail-case caddisflies (Trichoptera: Helicopsychidae) and new records of caddisflies from Chapada Diamantina, Bahia, Brazil. Zootaxa 4223: 343-358. https://doi.org/10.11646/zootaxa.4223.3.3.
https://doi.org/10.11646/zootaxa.4223.3.... , Vilarino & Calor 2017VILARINO A & CALOR AR. 2017. Trichoptera of Serra da Jibóia, Bahia, Brazil: new species of Helicopsyche (Helicopsychidae) and new records. Zootaxa 4311: 503-522. https://doi.org/10.11646/zootaxa.4311.4.4.
https://doi.org/10.11646/zootaxa.4311.4.... ), many of which have a restricted distribution in the NMAF (Table I).

Within this context, we can observe that the knowledge shortfalls related to species recognition and distribution (Linnean and Wallacean shortfalls) are accentuated mainly for the freshwater taxa, including Helicopsyche (Feropsyche) subgenus. Thus, here we describe and illustrate four new species of H. (Feropsyche) and provide new distribution records. In addition, we perform niche modeling based on the species distributions of the group to identify areas with high environmental suitability (high potential distribution) to direct future biodiversity research efforts on the NMAF, a highly endemic and underexplored ecoregion in Brazil.

MATERIALS AND METHODS

Study area

Abell et al. (2008)ABELL R ET AL. 2008. Freshwater ecoregions of the world: a new map of biogeographic units for freshwater biodiversity conservation. BioScience 58: 403-414. https://doi.org/10.1641/B580507.
https://doi.org/10.1641/B580507... classified the freshwater environments into ecoregions, among then it stands out the “Northeastern Mata Atlântica Freshwater” ecoregion (NMAF, number 328), which comprises all coastal drainages from the Sergipe River in the north to the Itabapoana river in the south. It is west bordered by the São Francisco Freshwater ecoregion in Northeast Brazil. NMAF comprises a mosaic of landscapes from mountains and valleys to sandstone plateaus with elevations from the flat coastal plain up to 2,890 m a.s.l. (Pico da Bandeira at Serra do Caparaó, the third highest peak of Brazil), and includes diverse phytophysionomies from the Atlantic Forest, Caatinga, and Cerrado domains (Hales & Petry 2013HALES J & PETRY P. 2013. Northeastern Mata Atlantica ecoregion. http://www.feow.org/ecoregions/details/northeastern_mata_atlantica. Accessed 08 September 2022.
http://www.feow.org/ecoregions/details/n... ).

Along with other freshwater environments, the drainages of the NMAF form a series of isolated hydrographic basins, which are separated by the scarped, mountainous landscapes of the eastern margin of the Brazilian crystalline shield (Ribeiro 2006RIBEIRO AC. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop Ichthyol 4: 225-246.). Resulting in a complex biogeographical history, with the hydrographic systems (e.g., Paraguaçú, Contas, Jequitinhonha, Doce, Paraíba do Sul), as well as several other more minor adjacent drainages, demonstrating a high rate of endemism (Ribeiro 2006RIBEIRO AC. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop Ichthyol 4: 225-246.).

NMAF is part of the 25 world hotspots of biodiversity, with highlighted importance for providing water and resource for the population, conservation of habitats, and maintenance of biodiversity (Myers et al. 2000MYERS N ET AL. 2000. Biodiversity hotspots for conservation priorities. Nature 403: 853-858. https://doi.org/10.1038/35002501.
https://doi.org/10.1038/35002501... ). Beside the Central Atlantic Forest Ecological Corridor (CAFEC), the ecoregion comprises part of the Chapada Diamantina Complex (CDC), and Serra do Espinhaço, recognized regions with environmental heterogeneity and high biodiversity (Santos et al. 2003SANTOS AMM, CAVALCANTI DR, SILVA JMC & TABARELLI M. 2003. Biogeographical relationships among tropical forests in northeastern Brazil. J Biogeo 34: 433-446., Silva & Castelletti 2005, Oliveira et al. 2015OLIVEIRA U, BRESCOVIT AD & SANTOS AJ. 2015. Delimiting Areas of Endemism through Kernel Interpolation. PLoS ONE 10: e0116633. https://doi.org/ 10.1331/journal.pone.0116633.
https://doi.org/ 10.1331/journal.pone.01... , Fricke et al. 2020FRICKE R, ESCHMEYER WN & VAN DER LAAN R. 2020. Eschmeyer’s catalog of fishes: genera, species, references. Academy of Science. San Francisco, California. http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain, Accessed 08 November 2022.
http://researcharchive.calacademy.org/re... , Santana et al. 2020SANTANA RO, DELGADO RC & SCHIAVETTI A. 2020. The past, present and future of vegetation in the Central Atlantic Forest Corridor, Brazil. Remote Sensing App: Soc Env 20: 100357. https://doi.org/10.1016/j.rsase.2020.100357.
https://doi.org/10.1016/j.rsase.2020.100... ), including high rates of endemism of several taxa (e.g., Camelier & Zanata 2014CAMELIER P & ZANATA AM. 2014. Biogeography of freshwater fishes from the Northeastern Mata Atlântica Freshwater ecoregion: distribution, endemism, and area relationships. Neot Ichthyol 12: 683-698. https://doi.org/10.1590/1982-0224-20130228.
https://doi.org/10.1590/1982-0224-201302... for fishes; Duarte et al. 2014DUARTE T, LECCI LS & CALOR AR. 2014. Stoneflies (Insecta: Plecoptera) from Serra Bonita, Bahia, Brazil: New species and updated records. Zootaxa 3779: 81-92. https://doi.org/10.11646/zootaxa.3779.1.9.
https://doi.org/10.11646/zootaxa.3779.1.... for stoneflies; Araujo et al. 2015 for beetles; Vilarino & Calor 2017VILARINO A & CALOR AR. 2017. Trichoptera of Serra da Jibóia, Bahia, Brazil: new species of Helicopsyche (Helicopsychidae) and new records. Zootaxa 4311: 503-522. https://doi.org/10.11646/zootaxa.4311.4.4.
https://doi.org/10.11646/zootaxa.4311.4.... for caddisflies; SOS Mata Atlântica & INPE 2017 for plants; Cavarzere et al. 2019CAVARZERE V, ALBANO C, TONETTI VR, PACHECO JF, WHITNEY BM & SILVEIRA LF. 2019. An overlooked hotspot for birds in the Atlantic Forest. Pap Avulsos Zool 59: e20195905. https://doi.org/10.11606/1807-0205/2019.59.05.
https://doi.org/10.11606/1807-0205/2019.... for birds).

Systematics

The material examined comprises 50 localities of the NMAF (Fig. 1a). Methods used in the preparation, examination, and illustration followed BlahnikBLAHNIK RJ, HOLZENTHAL RW & PRATHER AL. 2007. The lactic acid method for clearing Trichoptera genitalia. In: Bueno-Soria J, Barba-Alvarez R & Armitage BJ (Eds), Proceedings of the 12th International Symposium on Trichoptera. The Caddis Press, Columbus, Ohio, U.S.A., p. 9-14. et al. (2003) and Blahnik & Holzenthal (2004)BLAHNIK RJ & HOLZENTHAL RW. 2004. Collection and curation of Trichoptera, with an emphasis on pinned material. Nectopsyche, Neot Trichoptera Newsl, 1:8-20.. The terminology applied to the morphological structures followed Johanson (2002)JOHANSON KA. 2002. Systematic revision of American Helicopsyche of subgenus Feropsyche (Trichoptera: Helicopsychidae). Insects Syst Evol 60: 1-151., except for inferior appendage (rather than gonocoxite), as suggested by Holzenthal et al. (2016)HOLZENTHAL RW, BLAHNIK RJ & CALOR AR. 2016. Three new species of Helicopsyche von Siebold (Trichoptera: Helicopsychidae). from Brazil. Zootaxa 4038: 344-353. http://doi.org/10.11646/zootaxa.4038.1.29.
https://doi.org/10.11646/zootaxa.4038.1.... (Supplementary Material - Fig. S1).

The illustrations were made with a microscope equipped with a camera lucida, scanned, and finalized in Adobe® Illustrator® CS5. Microphotographs were made using a Leica stereoscope equipped with a digital camera, Nikon model DS-Fi1 and finalized in Adobe® Illustrator® CS5. Map was made using QGIS 3.10.10 (QGIS Development Team) and finalized with Adobe® Illustrator® CS5.

Type material will be deposited at Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZSP) and Museu de História Natural da Bahia, Bahia, Salvador, Brazil (UFBA), as indicated in the material examined. Additional material will be deposited at UFBA.

The distribution of caddisfly species through terrestrial ecoregions was used to estimate the number of unknown species in the NMAF using non-parametric estimators. Estimators were calculated based on incidence data (presence-absence only), using Brazilian phytogeographic domains as sampling unities, with the function specpool from the vegan package (Oksanen et al. 2019OKSANEN J ET AL. 2019. Vegan: Community Ecology Package. https://cran.r-project.org/web/packages/vegan.
https://cran.r-project.org/web/packages/... ) in software R. This function calculates three estimators: CHAO2, first-order jackknife (JACK1) and second order jackknife (JACK2). These non-parametric estimators of species richness help estimate a potential number of unobserved species based on incidence data as those available here. They have performed better than model-based or asymptotic estimators (Palmer 1990PALMER MW. 1990. The estimation of species richness by extrapolation. Ecology 71: 1195-1198. https://doi.org/10.2307/1937387.
https://doi.org/10.2307/1937387... , Hortal et al. 2006HORTAL J, BORGES PAV & GASPAR C. 2006. Evaluating the performance of species richness estimators: Sensitivity to sample grain size. J Animal Ecol 75: 274-287. http://doi.org/10.1111/j.1365-2656.2006.01048.x.
https://doi.org/10.1111/j.1365-2656.2006... ).

Niche modeling

Assumed there is a scarce number of records of species of the su1bgenus and that they present similar niches, areas, and feeding behavior. The species’ complete set of distributional records was used to understand the subgenus’s potential distribution in the NMAF. For this purpose, a database was compiled through the primary literature (species description and distributional records), Global Biodiversity Information Facility (GBIF; https://www.gbif.org), SpeciesLink (http://www.splink.org.br/), and original data from UFBA. Gazetteers and Google Maps© were used to register localization without coordinates. The centroid of the least comprehensive location was used. After the data compilation, a two steps filtering process was performed, (1) manual selection of the data with determined locality and species level; and (2) selection from the RStudio program (RStudio Team), discarding points that can generate an analysis bias (e.g., with equal coordinates or marine areas). After filtering, the database (Table SI) was used as input for niche modeling and to make a species distribution map.

Environmental data were obtained from monthly climate data for minimum, mean, and maximum temperature, precipitation, solar radiation, wind speed, water vapor pressure, and for total precipitation, 19 “bioclimatic” variables, and elevation on a scale of 30 arc seconds, available in the online database WorldClim version 2.1 (https://www.worldclim.org/data/worldclim21.html). After obtaining the data, in the RStudio program, a correlation analysis was performed using the Spearman method, the ‘correlate’ function of the ‘corrr’ package (Kuhn et al. 2022KUHN M, JACKSON S & CIMENTADA J. 2022. Package ‘corrr’: correlation in R. https://cran.r-project.org/web/packages/corrr.
https://cran.r-project.org/web/packages/... ). This procedure allows the selection of uncorrelated variables (correlation <30%) and avoids overweighing in the analyses.

Predictive distribution models are influenced by choice of modeling technique and the settings chosen by the researcher, summing up various uncertainties related to data quality and quantity, sample size, sampling bias, and spatial resolution (Zhang et al. 2015ZHANG L, LIU S, SUN P, WANG T, WANG G, ZHANG X & WANG L. 2015. Consensus forecasting of species distributions: The effects of niche model performance and niche properties. PloS one 10: e0120056. https://doi.org/10.1371/journal.pone.0120056.
https://doi.org/10.1371/journal.pone.012... ). To address these issues and improve distribution model performance, the use of an ensemble of algorithms, which address the results of multiple models in a single estimation, results in more accurate predictions than single model methods (Turner et al. 2018TURNER JA, BABCOCK RC, HOVEY R & KENDRICK GA. 2018. Can single classifiers be as useful as model ensembles to produce benthic seabed substratum maps? Estuar Coast Shelf Sc 204: 149-163. https://doi.org/10.1016/j.ecss.2018.02.028.
https://doi.org/10.1016/j.ecss.2018.02.0... ). Moreover, this methodology allows the identification of consensus forecasts by determining the level of agreement/disagreement between individual models, thus mapping model uncertainty (Araújo & New 2007ARAÚJO MB & NEW M. 2007. Ensemble forecasting of species distributions. Trends Ecol Evo 22: 42-47. https://doi.org/10.1016/j.tree.2006.09.010.
https://doi.org/10.1016/j.tree.2006.09.0... ).

Thus, we use at least one representative of the three main types of modeling algorithms to determine the level of agreement/disagreement between the different individual models building a more precise consensus. Four correlative modeling algorithms were used: two environmental distance models, the Bioclim (Nix 1986NIX HA. 1986. A biogeographic analysis of Australian elapid snakes. Atlas of Elapid Snakes of Australian. Atlas of Elapid Snakes of Australia. Canberra: Australian Gov. Publ. Service.) and Domain (Carpenter et al. 1993CARPENTER G, GILLISON AN & WINTER J. 1993. DOMAIN: a flexible modeling procedure for mapping potential distributions of plants and animals. Biodivers Cons 2: 663-680. http://doi.org/10.1003/BF00051966.
https://doi.org/10.1003/BF00051966... ); a regression-based model, the Generalized Linear Model (GLM; Nelder & Wedderburn 1932NELDER JA & WEDDERBURN RWM. 1932. Generalized linear models. J R Stat Soc Ser 135: 330-384.), and a machine learning model the Vector Support Machine (SVM; Tax & Duin 2004TAX D & DUIN R. 2004. Support Vector Data Description. Machine Learning 54: 45-66. http://doi.org/10.1023/B:MACH.0000008084.60811.49.
https://doi.org/10.1023/B:MACH.000000808... ).

To generate the absence points, not available for the species used, we randomly generated 72 pseudo-absence points (1:1 ratio for the occurrence points) through the ‘randomPoints’ function of the ‘dismo’ package (Hijmans et al. 2017HIJMANS RJ, PHILLIPS S, LEATHWICK J & ELITH J. 2017. Dismo package: Species distribution modeling. https://cran.r-project.org/web/packages/dismo.
https://cran.r-project.org/web/packages/... ) in the RStudio program. Data partitioning was randomly performed in 30% for training and 70% for testing the models. The repeatability of the models used to increase the robustness of the result was 200 times. Only models with Area Under Curve (Hanley & McNeil 1982HANLEY JA & MCNEIL BJ. 1982. The meaning and use of the area under a receiver operating characteristic ROC. curve. Radiology 143: 29-36. https://doi.org/10.1148/radiology.143.1.3063343.
https://doi.org/10.1148/radiology.143.1.... ) superior to 80% were considered for constructing suitability maps, using default limits of presence and absence.

RESULTS

Systematics

Family Helicopsychidae Ulmer, 1906

Genus Helicopsyche von Siebold, 1956

Subgenus Helicopsyche (Feropsyche) Johanson, 1998

Helicopsyche (Feropsyche) diamantina nov. sp.

Diagnosis. The new species is distinguished from all congeners by the following set of male characters: (i) abdominal segment X long and narrow, with convex lateral margins, and (ii) apex with a short, shallow cleft U-shaped, in dorsal view (Fig. 2j); (iii) inferior appendage with acuminated posterior projection, (iv) ventral margin with medial small setose projection, in lateral view (Fig. 2e), and (v) basomesal lobe of inferior appendage rounded, without a distinct protuberance or projection, in ventral view (Fig. 2f). The new species is morphologically similar to H. mateusi nov. sp., H. monda FlintFLINT OS JR. 1983. Studies of Neotropical Caddisflies, XXXIII: New Species from Austral South America (Trichoptera). Smithso Contrib Zool 377: 1-100., 1983 and H. obscura RuedaRUEDA-MARTÍN PA & ISA-MIRANDA AV. 2015. Association of immature stages of some caddisfly species from northwestern Argentina with description of a new species of Helicopsyche (Trichoptera: Helicopsychidae). Zootaxa 3949: 203-216. https://doi.org/10.11646/zootaxa.3949.2.3.
https://doi.org/10.11646/zootaxa.3949.2.... Martín & Isa Miranda 2015 by general shape of abdominal segment X in dorsal view, and basomesal lobe in ventral view, but differs from them by abdominal segment X with apex upcurved, in lateral view (Fig. 2e) (posteriorly oriented in all others), and apex with short, shallow apical cleft forming two lobes covered by setae, in dorsal view (Fig. 2j) (apical cleft absent in H. mateusi nov. sp. and H. obscura, and present, but without lobes in H. monda); The new species and H. monda share the inferior appendage medial region slightly constricted, in lateral view (wide in H. mateusi nov. sp. and H. obscura), but the new species differs from H. monda by the posteroventral margin of inferior appendage, new species with medial small setose projection versus without setose projection in H. monda, and by the basomesal lobe with posterior margin wide with rounded apex, in lateral view (Fig. 2e) (globose with short distal projection in H. mateusi nov. sp., subtriangular in H. monda, and unseen in H. obscura).

Description. Overall color yellowish brown (in alcohol, n=10). Forewing length 3.37–4.74 mm (n = 10), forks I, II, III and V present, with discoidal and thyridial cells, without medial cell. Hind wing forks I and V present, without discoidal and thyridial cells. Head: antennae around ≤1.4x forewing length, scape longer, length around half of head, covered by long setae (Fig. 2c); dense set of interantennal setae (Fig. 2c); postantennal warts finger-shaped with medial constriction with around 1/3 of head length; cephalic warts subtriangular covered by long setae (Fig. 2a); postocular warts subtriangular with wide base, covered by long setae (Fig. 2a); maxillary palps 2-segmented, covered by long setae (Fig. 2b); labial palps 3-segmented, covered by long setae (Fig. 2b). Thorax: pronotum bearing setal warts, digitated, with long and ferruginous setae; mesoscutum diamond-shaped, setal warts bean shaped covered by small setae, with 1/3 of mesoscutum length; mesoscutellum with setal warts subtriangular with small setae (Fig. 2d); legs with tibial spur formula 2, 2, 4. Abdomen: abdominal sternum VI process present, about 2/3 of the segment length (Fig. 2g).

Male genitalia. Abdominal segment IX anterior lobe rounded projection, located ventrally on segment, anterodorsal, and anteroventral margin concave, in lateral view (Fig. 2e); posterior lobe convex, basal plate V-shaped, in ventral view (Fig. 2f). Preanal appendages globose, in lateral view (Fig. 2e), digitated, short, in dorsal view (Fig. 2j). Abdominal segment X base wider than the apex; apex slightly rounded, upcurved, in lateral view (Fig. 2e); rectangular, with convex lateral margins, and apex with short, slightly deep apical cleft forming two lobes covered by setae, in dorsal view (Fig. 2j). Inferior appendage clavate, acuminated in anterior region, medial region slightly constricted, posterodorsal margin smooth and convex, posteroventral margin slightly concave with medial small setose projection, posterior region of appendage with acuminated projection, in lateral view (Fig. 2e); base wide, median constriction, apex wide, without apical tooth and inner face margin containing rows of long setae, in ventral view (Fig. 2f); basomesal lobe of inferior appendage globose covered by long setae, in lateral view (Fig. 2e), and wide rounded, slightly projected, covered by long setae, in ventral view (Fig. 2f). Phallus tubular, phallobase constricted medially (Figs. 2h,i), posterior region wide and rounded, in lateral view (Fig. 2h,i); phallotremal sclerite single, U-shaped, in ventral view (Fig. 2i).

Type material. Holotype: Brazil, Bahia, Complexo da Chapada Diamantina, Lençóis, Rio Mucugezinho, 12°23’44”S, 41°25’01”W, 306 m a.s.l., UV light pan trap, 1 ♂, 29.x.2013, Calor, Dias and Campos cols. (MZUSP). Paratypes: Brazil, Bahia, Complexo da Chapada Diamantina, Abaíra, Serra do Barbado, Tijuquinha abaixo, 13°11’56.3”S, 41°53’21.5”W, UV light pan trap, 2 ♂, 05.xi.2013, Calor, Dias and Campos cols. (MZUSP); same data, except 2 ♂, (UFBA); same data, except Igatú, Rio Coisa Boa, 12°53’23.3”S, 41°19’0.0”W, 633 m a.s.l., UV light pan trap, 1 ♂, 12.v.2010 (MZUSP); same data, except 9 ♂ (UFBA); same data, except Andaraí, Rio Piaba, 12°59’34”S, 41°20’23”W, 25♂, 22.i.2018, Calor et al. cols. (UFBA); same data, except Mucugê, Parque Municipal de Mucugê, Córrego Bandeira, 12°59’56.8”S, 41°19’53.8”W, 958 m a.s.l., UV light sheet attraction, 1 ♂, 01.vi.2019, Calor et al. cols. (UFBA); Córrego Boiadeiro, 2 ♂, 10.i.2015, Dias and Campos cols. (UFBA); same data, except, Rio Mucugê, 12°53’1.8’’S, 41°16’33’’W, 993 m a.s.l., UV light pan trap, 6 ♂, 25.xi.2018, Calor et al. cols. (MZUSP); same data, except, Rio Piabinha, 12°59’34”S, 41°20’23”W, 921 m a.s.l., UV light pan trap Branca/UV, 4 ♂, 25.vii.2010, Calor, Lecci Quinteiro, França, Arantes and Camelier cols. (UFBA); same data, except, Rio Tiburtino, 12°59’53’’S, 41°20’50’’W, 909 m a.s.l., UV light pan trap, 8 ♂, 13.v.2015, Calor et al. cols. (UFBA); same data, except Rio Cumbuca, 12°59’51”S, 41°20’56”W, UV light pan trap, 14 ♂, 23.vii.2010, Calor, Lecci Quinteiro, França, Arantes and Camelier cols. (UFBA); same data, except Palmeiras, Vale do Capão, riachinho (ponte), 12°34’19.2’’S, 41°30’52.5’’W, 918 m a.s.l., UV light pan trap, 1 ♂, 25.vi.2011, Calor, Camelier and Burguer cols. (UFBA); Piatã, Cachoeira do Patrício (embaixo), 13°05’12”S, 41°51’12”W, Light, 1 ♂, 29.x–03.xi.2013, Menezes cols. (MZUSP); Brazil, Bahia, Complexo da Chapada Diamantina, Pindobaçu, Cachoeira da Fumaça, 10°28’43”S, 40°12’23”W, 13 ♂, 13.xii.2009, Zacca, T. cols. (UFBA)

Distribution. NMAF (CDC) [Brazil (Bahia state)].

Remarks. This new species belongs to the Helicopsyche monda complex.

Etymology. The specific name, a noun in apposition, refers to the Chapada Diamantina Complex, a mountain range of the Caatinga domain and central region of Bahia State, Brazil, which is the type locality of new species. The specific name “diamantina” in Portuguese means “diamantiferous” and is an allusion to the large diamond reservoirs in the region.

Helicopsyche (Feropsyche) mateusi nov. sp.

(Fig. 3)

Diagnosis. The new species is distinguished from all congeners by the following set of male characters: (i) abdominal segment X quadrangular, with parallel lateral margins, (ii) apex slightly undulated not forming lobes, in dorsal view (Fig. 3j); (iii) inferior appendage with ventral margin substraight with mesal setose projections, and (iv) basomesal lobe globose with short distal projection, in lateral view (Fig. 3e); (v) inferior appendage with inner face bearing set of projections with spine-like setae, in ventral view (Fig. 3f). The new species is morphologically similar to Helicopsyche alajuela JohansonJOHANSON KA & HOLZENTHAL RW. 2010. The snail-case caddisfly subgenus Helicopsyche (Feropsyche) in Costa Rica, with the description of 3 new species (Trichoptera: Helicopsychidae). Zootaxa 2689: 37-47. https://doi.org/10.11646/zootaxa.2689.1.4.
https://doi.org/10.11646/zootaxa.2689.1.... & Holzenthal, 2010, H. diamantina nov. sp. and H. monda by general shape of inferior appendage in lateral view, and basomesal lobe in ventral view, but differs from them by the inner face of the inferior appendage bearing projections with spine-like setae (projections absent in all, except sometimes present in H. diamantina nov. sp.), and by the presence of basomesal lobe with posterior region wide, with acuminated apex, in lateral view (Fig. 3e) (absent in H. alajuela, and with rounded apex in H. diamantina nov. sp., with pointed apex in H. monda).

Description. Overall color yellowish brown (in alcohol). Forewing length 2.98–3.92 mm (n = 8), forks I, II, III, and V present, with discoidal and thyridial cells, without medial cell. Hind wing forks I and V present, without discoidal and thyridial cells. Head: antennae around ≤1.2x forewing length, scape longer, length subequal to head, with a dense set of interantennal setae (Fig. 3a); postantennal warts club-shaped with apex globose and around 1/3 of head length; cephalic warts subtriangular covered by long setae (Fig. 3a); postocular warts mid-moon shaped with wide base, covered by long setae (Fig. 3b); maxillary palps 2-segmented, covered by long setae (Fig. 3b); labial palps 3-segmented, covered by long setae (Fig. 3b). Thorax: pronotum bearing setal warts, digitated, with small setae; mesoscutum diamond-shaped, setal warts bean shaped covered by small setae, with 1/3 of mesoscutum length; mesoscutellum with setal warts subtriangular with small setae (Fig. 3c); legs with tibial spur formula 2, 2, 4. Abdomen: abdominal sternum VI process present, about half the segment length (Fig. 3g).

Male genitalia. Abdominal segment IX anterior lobe rounded projection, located midway on segment, anterodorsal margin sub-straight, anteroventral margin concave, in lateral view (Fig. 3e), posterior margin convex, basal plate V-shaped, in ventral view (Fig. 3f). Preanal appendages globose, in lateral view (Fig. 3e), digitated, in dorsal view (Fig. 3j). Abdominal segment X base wider than the apex, apex rounded, slightly upcurved, in lateral view (Fig. 3e); quadrangular, with parallel lateral margins, and apex slightly undulated, in dorsal view (Fig. 3j). Inferior appendage clavate, acuminated in anterior region, medial region wide; ventral margin substraight with medial setose projections, posterior region of appendage with finger shape projection, in lateral view (Fig. 3e); base wide, narrowing towards the acuminated apex and inner face margin of the inferior appendage containing projections with spine-like setae, in ventral view (Fig. 3f); basomesal lobe globose with short distal projection, in lateral view (Fig. 3e); wide, triangular, slightly projected, covered by long setae, in ventral view (Fig. 3f). Phallus tubular, phallobase constricted medially (Figs. 3h,i), acuminated in posteroventral region, slightly downcurved, in lateral view (Fig. 3h); phallotremal sclerite single, comma shaped, in ventral view (Fig. 3i).

Type material. Holotype: Brazil, Bahia, Complexo da Chapada Diamantina, Lençóis, Rio Mucugezinho, 12°23’44”S, 41°25’1”W, 306 m a.s.l, 1 ♂, 29.x.2013, UV light pan trap, Calor AR, Dias ES, and Campos R cols. (MZUSP). Paratypes. same data Holotype, except 3 ♂ (UFBA); same data, except 1 ♂ (MZUSP); same data, except 12°23’45”S, 41°24’56”W, 305 m a.s.l, 1 ♂, 01.viii.2010, UV light pan trap, Calor AR, Camelier P, Lecci L, Arantes T, and França D cols. (MZUSP); same data, except Complexo da Chapada Diamantina, Rio Ribeirão, 12°35’13.0’’S, 41°22’96.3’’W, 361 m a.s.l), 1 ♂, 23.x.2008, UV light pan trap, Calor AR, Mariano R, and Mateus S cols.; same data, except 1 ♂ (UFBA).

Distribution. NMAF (CDC) [Brazil (Bahia State)].

Remarks. This new species belongs to the Helicopsyche monda complex.

Etymology. The specific name is in honor of the eminent German scientist Dr. Sidnei Mateus (USP, Ribeirão Preto), an honorable citizen of Pedregulho municipality, for his friendship and contribution to several aquatic insects’ fieldwork.

Helicopsyche (Feropsyche) miltonsantosi nov. sp.

Diagnosis. The new species is distinguished from all congeners by the following set of male characters: (i) Abdominal segment X subquadrangular, with parallel lateral margins, and (ii) apex with shallow concavity, in dorsal view (Fig. 4j); and (iii) inferior appendage ventral margin strongly sinuous with medial setose projections, in lateral view (Fig. 4e); (iv) and with inner face bearing set of projections with spine-like setae, in ventral view (Fig. 4f); (v) basomesal lobe triangular, well projected, covered by long setae, in ventral view (Fig. 4f). The new species is morphologically similar to H. catoles Souza, Gomes & Calor, 2017, H. mateusi nov. sp. and H. paprockii JohansonJOHANSON KA. 1998. Phylogenetic and biogeographic analysis of the family Helicopsychidae (Insecta: Trichoptera). Entomol Scandinavica Suppl 53: 1-172. & Malm, 2006 by general shape of abdominal segment X, in dorsal view, and inferior appendage, in lateral view, but it differs from them by inferior appendage with basomesal lobe well projected, in ventral view (Fig. 4f) (slightly projected in H. catoles and H. paprockii, and not projected in H. mateusi nov. sp.). The new species and H. paprockii share the basomesal lobe of the inferior appendage triangular, in lateral view (Fig. 4e) (globose in H. catoles and H. mateusi nov. sp.), but the new species differs from the H. paprockii by the distal region of inferior appendage, in lateral view (Fig. 4e) (digitated in new species versus truncated in H. paprockii).

Description. Overall color yellowish brown (in alcohol). Forewing length 3.06–3.63 mm (n = 10), forks I, II, III, and V present, with discoidal and thyridial cells, without medial cell. Hind wing forks I and V present, without discoidal and thyridial cells. Head: antennae around ≤1.2x forewing length, scape longer, length subequal to head, with a set of interantennal setae (Fig. 4c); postantennal warts filiform around half of head length; cephalic warts globose covered by long setae (Fig. 4a); postocular warts mid-moon shaped with wide base, covered by long setae (Fig. 4b); maxillary palps 2-segmented, covered by long setae (Fig. 4b); labial palps 3-segmented, covered by small and ferruginous setae (Fig. 4b). Thorax: pronotum bearing setal warts, oval-shaped, with small setae; mesoscutum diamond-shaped, setal warts bean shaped covered by small setae, with 1/4 of mesoscutum length; mesoscutellum with setal warts subtriangular with small setae (Fig. 4d); legs with tibial spur formula 2, 2, 4. Abdomen: abdominal sternum VI process present, about half the segment length (Fig. 4g).

Male genitalia. Abdominal segment IX anterior margin with rounded projection, located ventrally on segment, anterodorsal, and anteroventral margin concave, in lateral view (Fig. 4e); posterior margin convex, basal plate V-shaped, in ventral view (Fig. 4f). Preanal appendages clavate, in lateral view (Fig. 4e), digitated, in dorsal view (Fig. 4j). Abdominal segment X base wider than the apex, without basal projection, apex truncated, without curvature, in lateral view (Fig. 4f), abdominal segment X subquadrangular, with parallel lateral margins and apex with shallow concavity, in dorsal view (Fig. 4j). Inferior appendage clavate, rounded in anterior region; medial region constricted and bent 90°; ventral margin strongly sinuous with medial setose projections, posterior region of appendage with short finger shape projection, in lateral view (Fig. 4e); base and apex with subequal width, apex without apical tooth, and inner face margin of the inferior appendage containing medial projections with spine-like setae in ventral view (Fig. 4f); basomesal lobe subtriangular, in lateral view (Fig. 4e); and triangular, well projected, covered by long setae, in ventral view (Fig. 4f). Phallus tubular, phallobase slightly constricted medially (Figs. 4h, i), posteroventral region acuminated, slightly downcurved, in lateral view (Fig. 4h); phallotremal sclerite single, U-shaped, in ventral view (Fig. 4i).

Type material. Holotype: Brazil, Bahia, Sebastião Laranjeiras, Riacho Mandiroba, 14°22’34.5”S, 43°02’18.9”W, 1 ♂, 5.v.2013, UV light pan trap, Nogueira M.A.M. col. (MZUSP). Paratypes: Brazil, Bahia, Sebastião Laranjeiras, Riacho Mandiroba, 14°22’34.5”S, 43°02’18.9”W, 8 ♂, 5.v.2013, UV light pan trap, Nogueira M.A.M. col. (UFBA); same data, except 6 ♂ (MZUSP), same data, except Abaíra, Distrito Catolés, 13°18’33.6”S, 41°51’62.9”W, 1263 m a.s.l., UV light pan trap, 1, 30.x.2013, Calor, Dias and Campos cols. (MZUSP); same data, except Piatã, Rio Toboró, 13°13’31”S, 41°44’43’’W, 860 m a.s.l., 1 ♂, 28.vii.2010, UV light pan trap, Calor, França, Quinteiro, Lecci, Camelier, and Arantes cols (UFBA).

Distribution. SFF and NMAFs (CDC) [Brazil (Bahia State)].

Etymology. This species is named in memory of Milton Almeida dos Santos (1926–2001), a Brazilian geographer, writer, scientist, journalist, lawyer, and university professor. Considered one of the most renowned intellectuals in Brazil in the twentieth century, he was one of the great names of the renovation of geography in Brazil that took place in the 1930s (Elias 2002ELIAS D. 2002. “Milton Santos: a construção da geografia cidadã”. In: El ciudadano, la globalización y la geografía. Homenaje a Milton Santos. Scripta Nova. Revista electrónica de geografía y ciencias sociales, Universidad de Barcelona.). Although he graduated in Law, he stood out for his works in several areas of geography, especially in studies of Third World urbanization and for his works on globalization in the 1990s (Elias 2002ELIAS D. 2002. “Milton Santos: a construção da geografia cidadã”. In: El ciudadano, la globalización y la geografía. Homenaje a Milton Santos. Scripta Nova. Revista electrónica de geografía y ciencias sociales, Universidad de Barcelona.). His work was characterized by a critical position on the capitalist system and its theoretical assumptions, dominant in the geography of his time (Elias 2002ELIAS D. 2002. “Milton Santos: a construção da geografia cidadã”. In: El ciudadano, la globalización y la geografía. Homenaje a Milton Santos. Scripta Nova. Revista electrónica de geografía y ciencias sociales, Universidad de Barcelona.). Thus, we used the specific name miltonsantosi as a tribute to the honorable citizen of the municipality of Brotas de Macaúbas in the Chapada Diamantina Complex region.

Helicopsyche (Feropsyche) paulofreirei nov. sp.

Diagnosis. The new species is distinguished from all congeners by the following set of male characters: (i) abdominal segment X trapezoid with convex lateral margins, (ii) apex with shallow concavity, not forming lobes, in dorsal view (Fig. 5j); (iii) inferior appendage subtriangular, medial region constricted; (iv) with posterior region wide, elongated with rounded apex, in lateral view (Fig. 5e); and (v) basomesal finger shaped, well developed, base 1.5x as wide as apex and apex covered by setae, in ventral view (Fig. 5f). The new species is morphologically similar to Helicopsyche cipoensis Johanson & Malm, 2006 and H. guara Holzenthal, Blahnik & Calor, 2016 by general shape of abdominal segment X and inferior appendage in lateral view, but differs from them by abdominal segment X trapezoid, apex with shallow concavity, not forming lobes, in dorsal view (Fig. 5j) (rectangular, with apex rounded in H. cipoensis, and subtriangular, with apex with very short, deep apical cleft forming two lobes covered by setae in H. guara); and basomesal lobe of inferior appendage base 1.5x as wide as apex, apex wide and truncated, in ventral view (Fig. 5f) (base and apex same width, apex wide and rounded in H. cipoensis and base 2x as wide as apex, apex narrow and finger shaped).

Description. Overall color yellowish brown (in alcohol). Forewing length 3.32–4.87 mm (n = 10), forks I, II, III, and V present, with discoidal and thyridial cells, without medial cell. Hind wing forks I and V present, without discoidal and thyridial cells. Head brownish; antennae around ≤1.2x forewing length, scape longer, length subequal to head, covered by long setae; with a dense set of interantennal setae (Fig. 5c); postantennal warts club-shaped, anterior region projected, posterior region wide and globose, around half of head length (Fig. 5a); cephalic warts subquadrangular margins with long setae (Fig. 5a); postocular warts subtriangular with wide base, covered by long setae (Fig. 5b); maxillary palps brown, 2-segmented, covered by long, ferruginous setae; labial palps brown, 3-segmented, covered by long, ferruginous setae (Fig. 5b). Thorax: pronotum brown, with setal warts, digitated, covered by long, ferruginous setae; mesoscutum brown, with setal warts, bean-shaped, covered by small, ferruginous setae; mesoscutellum brown, with setal warts, globose, covered by small, ferruginous setae (Fig. 5b); legs yellowish brown, tibial spur formula 2, 2, 4. Abdomen: yellowish brown; abdominal sternum VI process present, about two-thirds of the segment length (Fig. 5g).

Male genitalia. Abdominal segment IX anterior lobe rounded projection, located ventrally on segment, anterodorsal and anteroventral margins concave, in lateral view (Fig. 5e), posterior lobe convex, basal plate U-shaped, in ventral view (Fig. 5f). Preanal appendages globose, in lateral view, digitated, in dorsal view (Fig. 5j). Abdominal segment X base wider than the apex, apex pointed, in lateral view (Fig. 5e), lateral margin convex, with a pair of little apical projections, U-shaped apex cleft, weakly notches, with a row of setae near lateral margin and a set of setae at the apex, in dorsal view (Fig. 5j). Inferior appendage clavate, widest apically, rounded in posterior region, anterior margin convex, with margin smooth, ventral margin weakly sinuous, without notches, containing a row of setae near the margin, in lateral view (Fig. 5e); inner face margin of the inferior appendage covered with rows of long setae, base and apex with equal width, apex without apical tooth, in ventral view (Fig. 5f); basomesal lobe of inferior appendage digitated, well-developed, with posterodorsal region rounded, with a set of setae on the basomesal margin, a subapical spine-like setae and an apical set of short spine-like setae, in lateral view (Fig. 5e), digitated, covered of few short spine-like setae on margin, projected posteriad, with length 1/3 of inferior appendage, in ventral view (Fig. 5f). Phallus tubular, phallobase slightly constricted medially (Figs. 5h,i), acuminated in posteroventral region, downcurved, in lateral view (Fig. 5h); phallotremal sclerite single, U-shaped, in ventral view (Fig. 5i).

Type material. Holotype: Brazil, Bahia, Igrapiúna, Reserva Michelin, Trilha do Guigó, 1ª ponte, 13°49’21”S, 39°12’12”W, 1 ♂, 26.ix.2013, UV light pan trap 4, Equipe PARFOR (MZUSP). Paratypes: Brazil, Bahia, Igrapiúna, Reserva Michelin, Trilha do Guigó, 1ª ponte, 13°49’21”S, 39°12’12”W, UV light pan trap 4, 3 ♂, 26.ix.2013, Equipe PAFOR cols. (UFBA); same data, except, Córrego próximo ao alojamento, 13°49’23’’S, 39°10’21’’W, UV light pan trap, 1 ♂, 19.ix.2012, Equipe LEAq cols. (MZUSP); same data, except Córrego das Matas, Trilha do Guigó, 13°49’25.4”S, 39°12’10.8”W, 120 m a.s.l., 2 ♂, 22.ix.2012, UV light pan trap, Calor et al. cols. (UFBA)

Distribution. NMAF (CAFEC) [Brazil (Bahia State)].

Etymology. This species is named in memory of Paulo Reglus Neves Freire (1921–1993), a Brazilian educator and philosopher. He is considered one of the most remarkable thinkers in world pedagogy and the Patron of Brazilian Education (Ferreira & Wiggers 2018FERREIRA IV & WIGGERS ID. 2018. Brazilian Key-Thinkers on Education. Soc Register 22: 103-130.). His didactic practice was based on the premise that the student would assimilate the object of study by using a dialectic practice with reality (Ferreira & Wiggers 2018FERREIRA IV & WIGGERS ID. 2018. Brazilian Key-Thinkers on Education. Soc Register 22: 103-130.). Thus, we used the specific name paulofreirei as a tribute to all the educators who participated in fieldwork at the Reserva Ecológica Michelin during the biology undergraduate course in the context of Plano Nacional de Formação de Professores da Educação Básica (PARFOR).

New distributional records

Helicopsyche (Feropsyche) dinoprata DumasDUMAS LL & NESSIMIAN JL. 2019. New species of Helicopsyche von Siebold 1856 (Trichoptera: Helicopsychidae) from Brazil, including the redescription of Helicopsyche (Feropsyche) planorboides. Zootaxa 4619: 231-25. https://doi.org/10.11646/zootaxa.4619.2.2.
https://doi.org/10.11646/zootaxa.4619.2.... & Nessimian, 2019

[Type locality: Brazil, Rio de Janeiro, Santa Maria Madalena, Parque Estadual do Desengano, Morumbeca dos Marreiros, afluente do Ribeirão Macapá, 21°52’39.0” S, 41°54’55.3” W, 1,110 m; DZRJ; ♂].

Distribution. Northeastern Mata Atlântica, Paraíba do Sul, and Ribeira de Iguape Freshwater ecoregions [Brazil (Bahia and Rio de Janeiro states)].

Material examined. Brazil, Bahia, Amargosa, Faz. Sr. Alcides, Boqueirão, Colonha, 13°08’11”S, 39°39’46”W, 544 m, UV light pan trap, 1 ♂, 18.vii.2009, Calor and Lecci cols. (UFBA); same data, except Faz. Sr. Alcides, Boqueirão, Colonha, 13°08’11”S, 39°39’46”W, 544 m, UV light pan trap, 1 ♂, 18.vii.2009, Calor and Lecci cols. (UFBA); same data, except Camacan, Fazenda Waldemar da Farmácia, 15°25’13”S, 39°34’01”W, 310 m, UV light pan trap, 1 ♂, 28.iii.2011, Calor, Quinteiro, França and Barreto cols. (UFBA); same data, except, Wenceslau Guimarães, EEEWG, Riacho Dr. Germano, afluente Riacho Patioba, 13°34’50”S, 39°42’13”W, UV light pan trap, 1 ♂, 03.ix.2013, Calor, Duarte and Dias cols. (UFBA); same data, except Elísio Medrado, Serra da Jiboia, Reserva Jequitibá, 12°52’13’’S, 39°28’36.8’’W, 493 m, Malaise trap, 5 ♂, 05.iii.2013, Calor et al. cols. (UFBA)

Remarks. New record for the NMAF (CAFEC), as well as for the Brazilian Northeast region.

Helicopsyche (Feropsyche) guara Holzenthal, Blahnik & Calor, 2016

[Type locality: Brazil, Santa Catarina, [Blumenau] Rio Caeté, at the entrance to Parque Ecológico Spitzkopf, 23°00.350’S, 49°06.650’W, el. 92 m; MZUSP; ♂; ♀].

Distribution. São Francisco, Northeastern Mata Atlântica, and Southeastern Mata Atlântica Freshwater ecoregions [Brazil (Bahia, Minas Gerais, Santa Catarina, Rio de Janeiro states)].

Material examined. Brazil, Bahia, Cachoeira, Fazenda Villa Real, mata sede, 12°35’41”S, 38°53’58”W, 1 ♂, 15.vi.2003, Alvim, Souza, SilvaSILVA JMC & CASTELETI CHM. 2005. Estado da biodiversidade da Mata Atlântica brasileira. In: Galindo-Leal C & Câmara IG (Eds), Mata Atlântica: Biodiversidade, Ameaças e Perspectivas. Belo Horizonte, Fundação SOS Mata Atlântica-Conservação Internacional, p. 43-59., and Monteiro cols. (UFBA); same data, except Camacan, Fazenda Waldemar da Farmácia, 15°25’13”S, 39°34’01”W, 310 m a.s.l., UV light pan trap, 1 ♂, 28.iii.2011, Calor, Quinteiro, França and Barreto cols. (UFBA); same data, except UV light pan trap, 1 ♂, 28.iii.2011, Calor, Quinteiro, França and Barreto cols. (UFBA); same data, except Jandaíra, Reserva COPERN, 11°36’51.9’’S, 33°38’46.9’’W, UV light pan trap, 3 ♂, 11.x.2016, Kiszewski, Silva, Dias and Campos cols. (UFBA); same data, except Maracás, Milagres, MAMI 25AMA, 13°22’33.9”S, 40°29’22.0”W, 858 m a.s.l., UV light pan trap, 1 ♂, 25.iii.2012, PPBIO cols. (UFBA).

Remarks. New record for the NMAF (CAFEC), as well as for the Brazilian Northeast region.

Helicopsyche (Feropsyche) monda FlintFLINT OS JR. 1991. Studies of Neotropical Caddisflies, XLV: The Taxonomy, Phenology, and Faunistics of the Trichoptera of Antioquia, Colombia. Smithso Contrib Zool 520: 1-113., 1983

[Type locality: Paraguay, Depto. Alto Paraná, Salto del Monday, near Puerto Presidente Franco; NMNH; ♂].

Distribution. Orinoco Llanos, Northeastern Mata Atlântica, Paraíba do Sul, Lower Uruguay, Upper Uruguay, Upper Paraná and Lower Paraná Freshwater ecoregions [Argentina, Brazil (Bahia, Ceará, Minas Gerais, Paraná, Rio de Janeiro, Santa Catarina, São Paulo states), Paraguay, Venezuela].

Material examined. Brazil, Bahia, Camacan, RPPN Serra Bonita, córrego em frente ao barranco desmoronando, 15°23’13.6’’S, 39°33’56.3’’W, 333 m a.s.l., UV light pan trap, 2 ♂, 13.xi.2011, Quinteiro, Dias and Duarte cols. (UFBA); same data, except Córrego Itauna, 15°23’35”S, 39°33’50”W, 330 m a.s.l., UV light pan trap, 2 ♂, 29.iii.2011, Quinteiro, França and Barreto cols. (UFBA); same data, except córrego próximo ao alojamento, 13°49’24.6’’ S, 39°10’19.9’’ W, 63 m a.s.l., 2 ♂, 29.xii.2011, Quinteiro, Duarte and Dias cols. (UFBA); same data, except Complexo da Chapada Diamantina, Abaíra, Catolés de cima, Riacho da Forquilha (porteira), 13°13’28.3”S, 41°54’02.3”W, 1603 m a.s.l., 8 ♂, 02.xi.2013, Calor, Dias and Campos cols. (UFBA); same data, except Cachoeira do Guarda Mó, 13°19’35’’S, 41°19’35’’W, 126 m a.s.l., UV light pan trap, 1 ♂, 30.x.2013, Calor, Dias and Campos cols. (UFBA); same data, except Piatã, Rio de Contas, Cachoeira das Deusas, Fazenda Oshoki, 13°06’33.1”S, 41°50’20.6”W, UV light trap, 5 ♂, 05.xi.2013, Menezes col. (UFBA); same data, except Santa Teresinha, Pedra Branca, Serra da Jiboia, Riacho das Torres, 12°51’00”S, 39°28’48”W, 638 m a.s.l., UV light pan trap, 8 ♂, 24.xi.2010 (UFBA); same data, except Serra da Jiboia, 80 m abaixo da cachoeira, 12°51’00’’S, 39°28’48’’W, 638 m a.s.l., Malaise trap, 1 ♂, 08.viii–28.xi.2009, Calor and Lecci cols. (UFBA); same data, except Uruçuca, Serra Grande, Parque Estadual Serra do Conduru, Cachoeira da Trilha Principal, 14°29’48.5’’S, 39°03’53.1’’W, 223 m a.s.l., UV light pan trap ( branca), 3 ♂, 13.i.2014, Dias and Pereira cols. (UFBA).

Remarks. New record for NMAF (CAFEC and CDC), Brazil.

Helicopsyche (Feropsyche) petri Dumas & Nessimian, 2019

[Type locality: Brazil, Rio de Janeiro, Rio das Ostras, Reserva Biológica União, Trilha Interpretativa do Pilão, riacho, 22°25’29.2” S, 42°02’21.2” W; DZRJ; ♂].

Distribution. Northeastern Mata Atlântica, Paraíba do Sul, and Fluminense Freshwater ecoregions [Brazil (Bahia and Rio de Janeiro states)]

Material examined. Brazil, Bahia, Una, REBIO Una, 15°10’16.3”S, 39°03’40.5”W, 141 m a.s.l.) 2 ♂, 04.viii.2013, UV light pan trap, Dias, Campos, Laurindo, and Gudim cols. (UFBA)

Remarks. New record for NMAF (CAFEC), as well as for the Brazilian Northeast region.

Helicopsyche (Feropsyche) vergelana Ross, 1956

[Type locality: Mexico, Chiapas, Finca Vergel; INHS; ♂].

Distribution. Rio San Juan (Mexico), West Texas Gulf, Sorona, Rio Balsas, Papaloapan, Grijalva - Usumacinta, Chiapas - Fonseca, Quintana Roo - Motagua, Estero Real - Tempisque, San Juan (Nicaragua/Costa Rica), Chagres, Rio Tuira, North Andean Pacific Slopes - Rio Atrato, Magdalena - Sinu, South American Caribbean Drainages - Trinidad, Orinoco Piedmont, Orinoco Delta & Coastal Drainages, Essequibo, Guianas, Amazonas Guiana Shield, Madeira Brazilian Shield, Northeastern Caatinga & Coastal Drainages, Northeastern Mata Atlântica and Central Andean Pacific Slopes Freshwater ecoregions [Belize, Brazil (Bahia, Ceará, Maranhão, Pernambuco, Piauí states), Costa Rica, Grenada, Guatemala, Honduras, Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, Tobago, Trinidad, Venezuela].

Material examined. Brazil, Bahia, Iraquara, Pratinha, Rio Pratinha, abaixo da queda d’água, 12°21’10”S, 41°32’24”W, 642 m a.s.l., 2 ♂, 28.xi.2015, UV light pan trap, Queiroz, Santana, Mugnai, Ribeiro, Silva, and Cardoso cols. (UFBA)

Remarks. New record for NMAF (CDC), Brazil.

The four new species described here, and the new distribution records for five species (H. dinoplata, H. guara, H. monda, H. petri, and H. vergelana), increase from seven to 16 the known species in the NMAF. Our estimates indicate the existence of between 25 to 49 species of H. (Feropsyche) in the NMAF, based on the estimators of JACK1 (25 species), JACK2 (30 species), and CHAO2 (49 species).

Niche modeling

Sixteen species (twelve recorded and four new species) are used for modeling using distributional literature records, an online database, material examined, and data on UFBA (Table I). After correlation testing, seven raster variables were found to be uncorrelated, belonging to four groups, as arranged in Table II (more details Table SII and Fig. S2). Of the four algorithms tested, three (Bioclim, GLM, and SVM) presented AUC values higher than the cut-off value, and these were used to elaborate the subgenus environmental suitability maps (Fig. S3).

The distribution records of the present study are concentrated in the central and southern regions of the NMAF. Results point to high environmental suitability in two main areas of the NMAF region (Fig. 1b). The first area corresponds to the south of CDC, mainly in areas of high altitude (Fig. 1b), and the second is with high environmental suitability is the coastal drainages inserted in north of CAFEC (Fig. 1b). The other areas in the north and south of the NMAF show low environmental suitability (<0.5) (Fig. 1b).

DISCUSSION

Although H. (Feropsyche) is highly rich and virtually distributed in all freshwater ecosystems (Johanson 2002JOHANSON KA. 2002. Systematic revision of American Helicopsyche of subgenus Feropsyche (Trichoptera: Helicopsychidae). Insects Syst Evol 60: 1-151., Johanson & Malm 2006JOHANSON KA & MALM T. 2006. Seven new Helicopsyche (Feropsyche) Johanson, 2002 from the Neotropical region and Nearctic Mexico (Insecta: Trichoptera: Helicopsychidae). Zootaxa 1208: 1-24. https://doi.org/10.11646/zootaxa.1208.1.1.
https://doi.org/10.11646/zootaxa.1208.1.... ), its distributional range is poorly known, and its diversity may be underestimated, as evidenced by the richness estimates of the NMAF. The subgenus Feropsyche is usually sampled in small and medium-sized streams (Flint 1991). Despite the wide distribution and number of species in Brazil (23 species), only seven species of H. (Feropsyche) were known in the NMAF (Santos et al. 2020SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2020. Taxonomic Catalog of the Brazilian Fauna: order Trichoptera Insecta, diversity and distribution. Zoologia (Curitiba) 33: e46392. https://doi.org/10.3897/zoologia.37.e46392.
https://doi.org/10.3897/zoologia.37.e463... ), three of them known only from the type or adjacent locality. Despite slight differences, male genital structures have been remarkably useful for differentiating species in H. (Feropsyche). The shape and projections of the abdominal segment X, the inferior appendage, and the basomesal lobe have more useful characters in differentiating species. However, we observed differences in setal warts, especially of the head, when comparing species from different regions. These may indicate these as good characters for separating species groups in H. (Feropsyche).

Two species described here are part of what we designated, the Helicopsyche monda complex, a group of species with very similar genitalia morphology to H. monda. Helicopsyche monda was described by Flint (1983) from a series of specimens from Paraguay (holotype), Argentina, and Brazil (Santa Catarina state) (Flint, 1983), currently recorded from Northwestern (Oniroco Llanos Freshwater ecoregion, Venezuela) to Southeastern South American (Lower Uruguay Freshwater ecoregion, Argentina) (Holzenthal & Calor 2017HOLZENTHAL RW & CALOR AR. 2017. Catalog of the Neotropical Trichoptera (caddisflies). ZooKeys 654: 1-566. https://doi.org/10.3897%2Fzookeys.654.9516.
https://doi.org/10.3897%2Fzookeys.654.95... ). The original description by Flint (1983) shows divergence regarding the basomesal lobe of the inferior appendage when compared to the redescription and reillustration provided by Johanson (2002)JOHANSON KA. 2002. Systematic revision of American Helicopsyche of subgenus Feropsyche (Trichoptera: Helicopsychidae). Insects Syst Evol 60: 1-151.. In the ventral view, basomesal lobe of inferior appendage, according to Flint (1983), is slightly projected posteriad and subtriangular, although according to Johanson (2002)JOHANSON KA. 2002. Systematic revision of American Helicopsyche of subgenus Feropsyche (Trichoptera: Helicopsychidae). Insects Syst Evol 60: 1-151., the basomesal lobe is absent. However, no comment was made about the morphological differences.

Here we use Flint’s (1983) original description for comparison in differential diagnosis. However, all specimens morphologically identical to H. monda sensu Flint (1983) or Johanson (2002)JOHANSON KA. 2002. Systematic revision of American Helicopsyche of subgenus Feropsyche (Trichoptera: Helicopsychidae). Insects Syst Evol 60: 1-151. were designated as H. monda. These identifications indicate that what we know as H. monda is a complex of species, evidencing the need for a reanalysis of the type series and additional material to resolve this taxonomic problem.

Since the present work, 16 species have been recorded for the NMAF, including four new species and five new distribution records. Of these ten species are recorded from the Caatinga domain (only H. miltonsantosi does not occur in the CDC), six are recorded from the Atlantic Forest domain (all occurring in CAFEC and only H. guara and H. planorboides with records outside), and two are recorded from the Cerrado domain (H. angeloi and H. cipoensis) (Fig. 1a). The distribution records are mainly centered in easily accessible areas and protected areas, indicating that the same bias found for terrestrial organisms (i.e., Oliveira et al. 2016OLIVEIRA U ET AL. 2016. The strong influence of collection bias on biodiversity knowledge shortfalls of Brazilian terrestrial biodiversity. Divers Distrib 22: 1232-1244. https://doi.org/10.1111/ddi.12489.
https://doi.org/10.1111/ddi.12489... ), seems to apply for aquatic insects.

These results and richness estimates evidence the biodiversity deficits, especially Linnean and Wallacean shortfalls, in the NMAF and the Brazilian Northeast region. Given the vastness of the ecoregion, the limited number of distributional records and species reveal knowledge deficits in the north and south of the NMAF. Therefore, we pointed out two main areas, south of CDC and north of CAFEC (Fig. 1b), that have high environmental suitability (high potential distribution) and with piecemeal collection efforts in sparse areas, which highlights that these areas prioritized for research’s efforts as pointed out in results of the niche modeling.

The CDC presents a vegetation mosaic composed of Caatinga, Cerrado, and Campo Rupestres, among others (Velloso et al. 2002VELLOSO AL ET AL. 2002. Ecorregiões propostas para o bioma Caatinga. Associação Plantas do Nordeste. Instituto de Conservação Ambiental The Nature Conservancy do Brazil, 36 p.), and has essential springs of streams and rivers in the region (Juncá et al. 2005JUNCÁ FA, FUNCH L & ROCHA W. 2005. Biodiversidade e conservação da Chapada Diamantina. Brasília, Ministério do Meio Ambiente, Série Biodiversidade, 435 p.; Santos et al. 2020SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2020. Taxonomic Catalog of the Brazilian Fauna: order Trichoptera Insecta, diversity and distribution. Zoologia (Curitiba) 33: e46392. https://doi.org/10.3897/zoologia.37.e46392.
https://doi.org/10.3897/zoologia.37.e463... ). The region presents a high biodiversity of several groups, including Trichoptera. The CDC is where the most significant number of Trichoptera records are found in the NMAF, as well as the highest species richness and abundance (Santos et al. 2020SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2020. Taxonomic Catalog of the Brazilian Fauna: order Trichoptera Insecta, diversity and distribution. Zoologia (Curitiba) 33: e46392. https://doi.org/10.3897/zoologia.37.e46392.
https://doi.org/10.3897/zoologia.37.e463... ). CDC has received attention in the last decade (e.g., Research Program of Biodiversity in Semiarid region), resulting in many specimens housed at UFBA (and other biological collections in the Brazilian Northeast region). Besides, CDC presents several unexplored areas north and south of the ecoregion.

The second area of the NMAF with high environmental suitability is CAFEC, composed of a mosaic of Atlantic Forest with low- and high-altitude coastal regions (0 to 2868 m a.s.l.). Regarding land use, CAFEC presents ca. 41% occupied by forest, ca. 57% by some farming activity, and 2% formed by natural non-forest formation, non-vegetated areas, water bodies, and unobserved (Santana et al. 2020SANTANA RO, DELGADO RC & SCHIAVETTI A. 2020. The past, present and future of vegetation in the Central Atlantic Forest Corridor, Brazil. Remote Sensing App: Soc Env 20: 100357. https://doi.org/10.1016/j.rsase.2020.100357.
https://doi.org/10.1016/j.rsase.2020.100... ). The Atlantic Forest is the richest forest domain in Brazil, with 495 species of Trichoptera, of which 137 species are recorded for the NMAF (Santos et al. 2020SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2020. Taxonomic Catalog of the Brazilian Fauna: order Trichoptera Insecta, diversity and distribution. Zoologia (Curitiba) 33: e46392. https://doi.org/10.3897/zoologia.37.e46392.
https://doi.org/10.3897/zoologia.37.e463... ). However, despite being in the Atlantic Forest domain, CAFEC is mainly located in the Northeast region, with the most significant deficit of biodiversity knowledge and great potential to harbor new taxa (Santos et al. 2020SANTOS APM, DUMAS LL, HENRIQUES-OLIVEIRA AL, SOUZA WRM, CAMARGOS LM, CALOR AR & PES AMO. 2020. Taxonomic Catalog of the Brazilian Fauna: order Trichoptera Insecta, diversity and distribution. Zoologia (Curitiba) 33: e46392. https://doi.org/10.3897/zoologia.37.e46392.
https://doi.org/10.3897/zoologia.37.e463... ). Here are increased from three to nine known species, but according to estimates and models presented, there is still much to be known, especially in the north of CAFEC, which has a larger area of forest cover and more outstanding environmental suitability for H. (Feropsyche).

Despite our work’s contributions that increase the known species from seven to 16, distribution records are still concentrated and scarce for most of the NMAF. Thus, we consider that this ecoregion should be prioritized for research efforts given the high environmental suitability (high potential distribution) for H. (Feropsyche) species and the diversity estimates that indicate that there is still much to be recorded and between 36–67% of species awaiting description. As priority areas within the ecoregion, we highlight the CDC and CAFEC.

CONCLUSIONS

This work is the first study to use distribution modeling tools for the order Trichoptera in the Neotropics. It provides essential data that help to reduce the knowledge of biodiversity shortfalls. It also points to ways that can help select areas for research efforts in the NMAF. Our data, although important, are still the first step in the direction of understanding the biodiversity of the ecoregion, so future studies and databases can provide data that help in a better understanding of the distribution of H. (Feropsyche), as well as a better resolution of the potential distribution models of species.

The main conclusions of the paper are (i) the need for collection and analysis of material in areas of high environmental suitability and without or scarce distributional records, optimizing the use of resources in taxonomic research; and (ii) the need for a detailed reanalysis of what becomes known as the Helicopsyche monda complex, to face the Linnean shortfall and solve this taxonomic problem.

ACKNOWLEDGMENTS

R. Pereira, also thanks to the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES, finance code 001, PDS-CAPES-88882.453922/2019-01) for the doctoral fellowship. 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 also thank Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) for the collection license. We also thank Michelin, especially Dr. Kevin Flesher and André Santos, for all their support at Reserve Ecológica Michellin. We thank the collectors and local guides for all support and material examined here.

SUPPLEMENTARY MATERIAL

Figure S1-S3.

Table SI-SII.

REFERENCES

Publication Dates

History