Hidden biodiversity in Neotropical streams: DNA barcoding uncovers high endemicity of freshwater macroinvertebrates at small spatial scales

6. Ramı´rez A, Paulson DR, Esquivel C. Odonata of Costa Rica: Diversity and checklist of species. Revista de Biologı´a Tropical. 2000; 48: 247–254. 7. Posada-Garcı´a JA, Rolda´n-Pe´ rez G. Clave ilustrada y diversidad de las larvas de trichoptera en el nor- occidente de Colombia / Illustrated key for the larvae of Trichoptera in the northwest of Colombia. Cal- dasia. 2003; 25: 169–192. 8. Springer M. Clave taxono´mica para larvas de las familias del orden Trichoptera (Insecta) de Costa Rica. 1. 2006; 273–286. 9. Flowers RW. Diversity of stream-living insects in northwestern Panama. J North Am Benthol Soc. 1991; 10: 322–334. 10. Quintero D, Aiello A. Insects of Panama and Mesoamerica: Selected Studies. Oxford University Press; 1992. 11. Melo AS, Froehlich CG. Macroinvertebrates in neotropical streams: richness patterns along a catch- ment and assemblage structure between 2 seasons. Journal of the North American Benthological Soci- ety. 2001. pp. 1–16. https://doi.org/10.2307/1468184 12. Boyero L, Bailey RC. Organization of macroinvertebrate communities at a hierarchy of spatial scales in a tropical stream. Hydrobiologia. 2001; 464: 219–225. 13. Holzenthal RW, Blahnik RJ, Prather AL, Kjer KM. Order Trichoptera Kirby, 1813 (Insecta), Caddisflies. 2007. Available: https://conservancy.umn.edu/handle/11299/196322 14. Ligeiro R, Melo AS, Callisto M. Spatial scale and the diversity of macroinvertebrates in a Neotropical catchment. Freshw Biol. 2010; 55: 424–435. 15. Merritt RW, Cummins KW. An introduction to the aquatic insects of North America. Kendall Hunt; 1996.

16. Jacobsen D, Cressa C, Mathooko JM, Dudgeon D. 4—Macroinvertebrates: Composition, Life Histories and Production. In: Dudgeon D, editor. Tropical Stream Ecology. London: Academic Press; 2008. pp. 65–105. 17. Fischer AG. Latitudinal variations in organic diversity. Evolution. 1960; 14: 64–81. 18. Rosenzweig ML. Species Diversity in Space and Time. Cambridge University Press; 1995. 19. Jacobsen D, Schultz R, Encalada A. Structure and diversity of stream invertebrate assemblages: the influence of temperature with altitude and latitude. Freshw Biol. 1997; 38: 247–261. 20. Boyero L. Short Communication: Insect biodiversity in freshwater ecosystems: is there any latitudinal gradient? Mar Freshwater Res. 2002; 53: 753–755. 21. Wantzen KM, Ramı´rez A, Winemiller KO. New vistas in Neotropical stream ecology—Preface. J North Am Benthol Soc. 2006; 25: 61–65. 22. Boyero L, Ramı´rez A, Dudgeon D, Pearson RG. Are tropical streams really different? J North Am Benthol Soc. 2009; 28: 397–403. 23. Mayhew PJ. Why are there so many insect species? Perspectives from fossils and phylogenies. Biol Rev Camb Philos Soc. 2007; 82: 425–454. https://doi.org/10.1111/j.1469-185X.2007.00018.x PMID: 17624962 24. Dijkstra K-DB, Monaghan MT, Pauls SU. Freshwater biodiversity and aquatic insect diversification. Annu Rev Entomol. 2014; 59: 143–163. https://doi.org/10.1146/annurev-ento-011613-161958 PMID: 24160433 25. Mu´rria C, Rugenski AT, Whiles MR, Vogler AP. Long-term isolation and endemicity of Neotropical

aquatic insects limit the community responses to recent amphibian decline. Diversity and Distributions. 2015. pp. 938–949. https://doi.org/10.1111/ddi.12343 26. Smith SA, Bermingham E. The biogeography of lower Mesoamerican freshwater fishes. J Biogeogr. 2005; 32: 1835–1854. 27. Aguilar C, Miller MJ, Loaiza JR, Gonza´ lez R, Krahe R, De Leo´n LF. Tempo and mode of allopatric diver- gence in the weakly electric fish Sternopygus dariensis in the Isthmus of Panama. Sci Rep. 2019; 9: 18828. https://doi.org/10.1038/s41598-019-55336-y PMID: 31827183 28. Monaghan MT, Wild R, Elliot M, Fujisawa T, Balke M, Inward DJG, et al. Accelerated species inventory on Madagascar using coalescent-based models of species delineation. Syst Biol. 2009; 58: 298–311. https://doi.org/10.1093/sysbio/syp027 PMID: 20525585 29. Garcı´a-Lo´ pez A, Mico´ E, Mu´ rria C, Galante E, Vogler AP. Beta diversity at multiple hierarchical levels: explaining the high diversity of scarab beetles in tropical montane forests. J Biogeogr. 2013; 40: 2134– 2145. 30. Dudgeon D, Arthington AH, Gessner MO, Kawabata Z-I, Knowler DJ, Le´vêque C, et al. Freshwater bio- diversity: importance, threats, status and conservation challenges. Biol Rev Camb Philos Soc. 2006; 81: 163–182. https://doi.org/10.1017/S1464793105006950 PMID: 16336747 PLOS ONE Hidden biodiversity in Neotropical streams PLOS ONE | https://doi.org/10.1371/journal.pone.0231683 August 7, 2020 11 / 13