Micro-habitat use and seasonality of caddisfly larvae (Trichoptera) in two streams in eastern Cuba

333 Micro-habitat use and seasonality of caddisfly larvae (Trichoptera) in two streams in eastern Cuba Pedro López Del Castillo1, Germán M. López Iborra2, Liliana María Gómez Luna3, Perla Alonso Eguía-Lis4 1 Centro Oriental de Ecosistemas y Biodiversidad (BIOECO), Enramada # 601, Santiago de Cuba, Cuba 2 Departamento de Ecología, IMEM Ramon Margalef, Universidad de Alicante, San Vicent del Raspeig, Spain 3 Centro Nacional de Electromagnetismo Aplicado, Universidad de Oriente, Ave. de las Américas s/n Esq. I Reparto Sueño CP 90400, Santiago de Cuba, Cuba 4 Instituto Mexicano de Tecnología del Agua, Blvd. Paseo Cuauhnáhuac 8532, Progreso, 62550 Jiutepec, Morelos, Mexico Corresponding author: Pedro López Del Castillo ([email protected]) Copyright: © Pedro López Del Castillo et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Four microhabitats within pools (cobbles, sand, leaf litter, and bank vegetation) and one microhabitat in riffles (cobbles) were sampled in both rainy and dry seasons to identify groups of caddisfly species that share larval microhabitats across 15 sites in streams in eastern Cuba. A total of 4,367 individuals representing 13 families, 22 genera, and 36 taxa (species and morphospecies) were collected. To explore the distribution of caddisfly species by microhabitat, a k-means clustering method was used. This analysis grouped the samples into seven clusters based on species abundance across microhabitats, seasonality, altitude, and stream order. The results show that caddisfly abundance is strongly influenced by microhabitat type, seasonal variation, and river characteristics. Riffles consistently had the highest abundance of caddisflies across all sampling periods. Functional feeding groups showed variable abundance and behavior. Scrapers were strongly influenced by seasonality with high abundance in the rainy season, while generalists showed no significant seasonal effect on abundance. This study provides detailed biological trait information for specific caddisfly species, which is essential for using multimetric indices to accurately determine the health of freshwater ecosystems. Key words: Assemblage composition, heterogeneity, k-means, tropical streams Introduction Caddisfly larvae are widely distributed in freshwater ecosystems, particularly in streams and rivers, where they play a crucial role in ecosystem functioning and offer several ecosystem services (Morse et al. 2019). The use of different habitats by caddisfly larvae has been widely studied (Ward 1992; Allan 1995; Urbanič et al. 2005; Martini and Waringer 2021). This group of freshwater insects exhibits diverse microhabitat use, which is influenced by ecological factors and their functional roles within aquatic ecosystems (Urbanič et al. 2005). Microhabitat utilization by larvae is driven mainly by substrate type, water flow, and the availability of resources, which in turn affect their distribution and abundance across different stream sections (Statzner et al. 2005). Academic editor: Ralph W. Holzenthal Received: 16 February 2025 Accepted: 2 July 2025 Published: 10 December 2025 ZooBank: https://zoobank.org/ F4616739-639B-41CC-B6778D0992FCB94A Citation: López Del Castillo P, López Iborra GM, Gómez Luna LM, Alonso Eguía-Lis P (2025) Micro-habitat use and seasonality of caddisfly larvae (Trichoptera) in two streams in eastern Cuba. In: Ríos-Touma B, Frandsen PB, Holzenthal RW, Houghton DC, Rázuri-Gonzales E, Pauls SU (Eds) Proceedings of the 18th International Symposium on Trichoptera. ZooKeys 1263: 333–350. https://doi.org/10.3897/ zookeys.1263.150346 ZooKeys 1263: 333–350 (2025) DOI: 10.3897/zookeys.1263.150346 334 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba Additionally, competition for food and space, predation, and prey distribution may also influence the abundance and distribution of aquatic insects (Minshall 1984). In streams, where conditions such as temperature, flow, and substrate types can vary greatly, caddisfly larvae exhibit specific behaviors in selecting microhabitats that help them optimize their survival and development (Poff and Ward 1989). The substrate is an important factor in determining habitat availability for caddisfly larvae. They can be found on rocks, woody debris, leaf litter, and submerged plants (Martini and Waringer 2021). Several key drivers, including hydrological connectivity, physicochemical factors, and life history traits influence the seasonal variation in caddisfly microhabitat use in the Neotropics. These factors interact, shaping the distribution and abundance of caddisfly larvae across different seasons (Van den Brink et al. 2013). These preferences are crucial for understanding their ecological impact and their role as bioindicators in stream environments (Brand et al. 2012). In the Caribbean Region, the studies conducted to date have explored a range of topics, including taxonomy, natural history, distribution, and abundance of freshwater macroinvertebrates (Alonso-EguíaLis et al. 2014; Ramírez and Gutiérrez-Fonseca 2014; Bastardo and Sánchez Rosario 2017; López Del Castillo et al. 2024). However, the information about the microhabitat use, selection, and preference of microhabitats at the species level of caddisflies is sparse and scattered since this information is primarily provided in ecological remarks to the species description (Botosaneanu 1994, 2002; Flint 1996; Naranjo López and González Lazo 2005). Therefore, the main goals of this study are to describe the microhabitats used by caddisfly larvae in two mountain rivers in eastern Cuba and to identify groups of caddisfly species that share the same microhabitats. Considering the influence of seasonal changes in water flow on microhabitat availability, analyses of seasonal variation in microhabitat use have also been conducted. Materials and methods Study area This study was carried out in the Yara and Nagua river basins on the north slope of Sierra Maestra Mountain System, approximately 40 km southwest of Bayamo city and bounded within 19°00'00"N and 20°09'17"N and 76°58'30"W and 76°51'23"W. Both watersheds are adjacent in the headwaters, and there is a convergency of the watercourse in Paso Malo water reservoir (Fig. 1). The geology of the watersheds is characterized by the undifferentiated presence of the El Cobre group, related to the Paleocene–Eocene period. Abundant rocks include andesites and tuffs, mixed with volcanic breccias (Núñez et al. 1989; Viña Bayés 2001). Also, ferralitic-lixiviated soil, brown without carbonates, and skeletal soils are abundant in the area (Hernández et al. 1999). The Yara headwater is located on the northern slope of Turquino National Park. It is characterized by a small portion of tropical cloud forest, small patches of coffee plantations, and other crop areas. The predominant riparian vegetation in the Yara River basin includes montane rainforest and mesophyll evergreen forest. On the other hand, the Nagua sub-watershed has some mesophyll evergreen forest, but other types of land use are present, such as intensive agriculture areas and coffee plantations. 335 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba The Cuban archipelago experiences two distinct weather seasons each year. The period from November to April is known as the “dry season,” characterized by lower rainfall. In contrast, the “rainy season” takes place from May to October (Ortiz and Rivero 2004; Allen and Mapes 2017). The annual precipitation in the study areas ranges from 1,800 to 2,200 mm (Puentes 2001). Sampling Collecting techniques and microhabitats Fifteen sampling sites were established in streams from second to fourth order (Strahler 1957) in the Yara (eight sites) and Nagua rivers (seven sites). The altitude ranged from 150 to 575 m above sea level, whereas the areas of the Figure 1. The drainage area of the Yara and Nagua river system and distribution of the sampling sites (n = 15). 336 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba sub-watersheds were between 0.3 and 98 km2 (Table 1). In the Yara River, four sites were distributed in streams of second order and the remaining four between third and fourth order. Seven points along the Nagua River were classified as thirdor fourth-order streams. The only exception was Las Cuevas site, which was classified as a second-order stream (Fig. 1). The seasonality effect was accounted for by sampling in two time periods representing different river flow conditions. Two samplings were carried out in March and April 2010, which corresponds to the end of the dry season, when the river flow is at its lowest. Additional sampling was conducted in November 2010 at the onset of the dry season. However, water levels remained high during that month in the study year, as October was the month with the highest rainfall and the highest number of rainy days (INRH 2011, Fig. 2). Table 1. Main features and location of sampling sites. Basins Sample sites Latitude, Longitude Stream order Altitude (m) Drainage area (km2) Yara Brazo Derecho 20°01'22"N, 76°50'49'′W 2594 2.7 Yara Brazo Izquierdo 20°02'08"N, 76°50'46'′W 2558 2.5 Yara La Jeringa 20°01'54'′N, 76°51'45'′W 3394 8.6 Yara San Francisco 20°02'02'′N, 76°51'53'′W 2517 2.7 Yara El Mogo 20°01'51'′N, 76°53'29'′W 2 637 0.3 Yara Santo Domingo 20°02'28'′N, 76°54'09'′W 4268 26.1 Yara Providencia 20°04'54'′N, 76°55'57'′W 4 116 75.2 Yara Palma Criolla 20°05'15'′N, 76°55'33'′W 3 134 14.7 Nagua Los Lirios 20°06'11'′N, 76°50'14'′W 3 266 10.7 Nagua Rancho Claro 20°05'48'′N, 76°50'39'′W 3259 22.7 Nagua Los Lajales 20°06'31'′N, 76°50'51'′W 4 231 36.6 Nagua Frio Nagua 20°06'45'′N, 76°52'11'′W 4168 55.1 Nagua Guasimilla 20°07'18'′N, 76°52'01'′W 3178 17.6 Nagua Las Cuevas 20°06'33'′N, 76°52'33'′W 2187 4.6 Nagua Sierrita de Nagua 20°07'44'′N, 76°55'13'′W 496 98.7 Figure 2. Total monthly rainy days (RD, columns, left axis) and precipitation (PP, lines, right axis) in 2010 in Yara and Nagua watersheds. 337 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba The sampling design included five microhabitats within two identified subsystems: pools and riffles. Microhabitat characterization followed the Wentworth grade scale, with modifications by Cummins (1962) and Vilenica et al. (2018), and considered substrate heterogeneity and depth, as per established methodologies (Coleman and Hynes 1970; Poole and Stewart 1976; Dudgeon 1982). Pool subsystems were the most heterogeneous, with four microhabitats—cobbles, sand, leaf litter, and riparian vegetation—present at all sites. In the riffle subsystem, cobbles were the only microhabitat present at all sampling sites, while other microhabitats were limited and poorly represented. Sampling in the pools was carried out using four methods, one for each microhabitat: a) direct picking-up of 15 cobbles (stones) from pools with no more than 20 cm depth, b) sand was sampled using a homemade D-net (0.5 mm mesh size) by shaking the bottom in an area of 30 × 30 cm, at 20 cm depth, c) for leaf litter, a pack of 30 × 30 cm, at 10 cm deep was collected d) bank vegetation was sampled by shaking the D-net inside the submerged bank vegetation or roots systems included in a length of 3 m along the bank. The samplings in the cobbles in riffles were made in the shallow “erosional zone”, where waters have a high flow speed, thus two complementary methods were used in this microhabitat to compensate for the loss of individuals that could be dragged by the current: a) direct picking-up out of cobbles; taking 15 cobbles with 10 cm of similar size and shape, as we did in pools b) D-Net was used to collect the caddisflies dragged by the flow when we agitated the cobbles in an area of 30 × 30 cm. Taxonomic identification All samples were preserved in ethyl alcohol at 90% during the sampling until identification in the laboratory. Organisms were sorted and counted. Identification of caddisfly larvae was made to species level with a stereomicroscope using the following keys and taxonomical description criteria: Botosaneanu and Sykora (1973), Botosaneanu (1977, 1993, 1994, 2002), Kumanski (1987), Flint (1996), Wiggins (1996, 2007). The morphospecies name corresponds to the description of Botosaneanu (1994). The specimens were preserved in 70% alcohol and stored at BIOECO’s Animal Biology Department collection after identification. Assignment of functional feeding groups Specimens were assigned to functional feeding groups (FFG) based on Merritt and Cummins (1996), Cummins et al. (2005), and Ramírez and Gutiérrez Fonseca (2014) using the genus or family level to each morphospecies. The defined groups were Predators (Pr), Scrapers (Sc), Shredders (Sh-Dt shredder on plant detritus) (Sh-Hb shredder on live plant tissue), Filters (Ft), Piercers (Pc-Hb), and Collectors-Gatherers (CG). Data analysis K-means clustering was used to identify groups of caddisfly species with similar use of microhabitats and seasonality (Legendre and Legendre 2012; Borcard et al. 2018). A matrix containing the total number of individuals of each species at 338 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba each microhabitat in the dry (the two sampling dates averaged) and rainy seasons (20 species × 10 microhabitats-season) was built. Following Borcard et al. (2018), this matrix was transformed using the Hellinger transformation and standardized, then a k-means partitioning analysis was performed to identify the optimal number of species groups, of species according to their use of microhabitats in both seasons. The Calinski criterion was used to identify the optimal number of groups (Borcard et al. 2018). These analyses were performed with the vegan package (Oksanen et al. 2011) in R v. 4.0.3 (R Core Team 2020). Results A total of 4,367 individuals were collected during the study, representing 13 families, 22 genera, and 36 infrageneric taxa (species and morphospecies). Of these, eight morphospecies with fewer than six specimens were excluded from the abundance analysis. This number represented 34% of the total morphospecies. Among these species were two morphospecies from the family Hydroptilidae (Orthotrichia sp. 1 and Neotrichia sp .1), and two from Helicopsychidae (Helicopsyche cubana (Kingsolver, 1964) and H. sp. (de)). Three families were represented by one morphospecies each: Austrotinodes cubanus (Kumanski, 1987) from Ecnomidae, Chimarra sp. 1 from Philopotamidae, and Macronema sp. 1 from Hydropsychidae. The k-means analysis revealed that the caddisfly samples clustered into seven distinct groups of species (Fig. 3) based on their abundance across microhabitats and seasonal patterns (Table 2). Based on the classification of functional feeding groups (FFG) by Ramírez and Gutiérrez Fonseca (2014), the two most diverse groups were scrapers and piercers with seven species each (Table 2). In addition, scrapers had the highest abundance (1,894), followed by filterers of the family Hydroptilidae (1,159). Figure 3. K-means cascade plot showing groups of caddisflies’ species in Yara and Nagua rivers. The left plot illustrates how the species are clustered across different k-means partitions, with the y-axis representing the number of groups in each partition and the x-axis representing the species and morphospecies. Each color denotes a distinct cluster within a given partition. The Calinski criterion posits that elevated values are indicative of more distinct clusters, and the red dot signifies the optimal cluster count. However, given the formation of single-species clusters, the number of clusters should be reduced to seven. 339 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba Group (I) (Fig. 4A) was integrated by the morphospecies Polycentropus sp. 1 and Helicopsyche near comosa which belong to the families Polycentropodidae and Helicopsychidae, respectively. This group reported the lowest abundance values (n = 21), with no individuals found in the rainy season. Polycentropus sp. 1 is a predator that uses all microhabitats except for the riffles. The sand microhabitat was the least used, pool stones, bank vegetation, and fallen leaves were also used with a similar proportion. On the other hand, Helicopsyche near comosa is a scraper that uses all microhabitats except sand. It is most frequently found in the bank vegetation microhabitat, followed by the riffles, among fallen leaves, and in pool stones (the last two less frequently). Group (II) (Fig. 4B) included two species, Marilia scudderi (Banks 1924) (Odontoceridae) and Helicopsyche sp. 1 (Helicopsychidae). During the rainy season, both species presented the highest abundance values, also a generalist behavior in the use of the microhabitats. Most often, pool stones and sand were used, followed by riffles. In contrast, the microhabitats of fallen Table 2. Grouping of species according to k-means analysis. The abundance column represents the mean number of individuals collected in the dry season and the rainy season samplings. The dominant microhabitat column identifies the microhabitat with the highest percentage of individuals averaged among species of each group in each season. Groups Abbreviation – Morphospecies – Functional Feeding Group (FFG). FFG: Pr = Predators, Sc = Scrapers, Sh = Shredders, (Sh-Dt shredder on plant detritus) (Sh-Hb Sh-Hb shredder on live plant tissue), Ft = Filters, Pc-Hb = Piercers, CG = Collectors-Gatherers. Abundance Dominant microhabitat use Dry Rainy Total I 1Pol_sp1. Polycentropus sp. 1 (Pr) 8 0 21 Generalist in dry season 2Hel_nea. Helicopsyche near comosa (Sc) 13 0 II 3Mar_scu. Marilia scudderi Banks, 1924 (Sh) 25 4204 Generalist in dry season 4Hel_sp1. Helicopsyche sp. 1 (Sc) 147 28 III 5Hyd_sp1. Hydropsyche sp.1 (Ft. Some Pr and seasonal Sc) 12 1 168 Pool stone and riffle in dry season 6Oxy_sp1. Oxyethira sp. 1 (Pc-Hb, Sc, CG) 78 3 7Och_sp1. Ochrotrichia sp. 1 (Pc-Hb, Sc, CG) 70 8Xip_cub. Xiphocentron cubanum Botosaneanu, 1993 (CG) 34 12 9Ant_sp1. Antillopsyche sp. 1 (Pr) 11 2 10Hel_cf. Helicopsyche cf. hageni (Sc) 9 0 IV 11Cer_sp1. Cernotina sp1 (Pr) 10 18 423 Generalist 12Phy_cha. Phylloicus chalybeus (Hagen, 1861) (Sh-Dt, Sc) 62 58 13Nec_cub. Nectopsyche cubana (Banks, 1938) (Sh-Hb, CG) 60 36 14Hel_hag. Helicopsyche hageni Banks, 1938 (Sc) 30 66 V 15Car_sp1. Cariboptila sp. 1 (Sc) 20 88 121 Pool stones in rainy season 16Hel_sp2. Helicopsyche sp. 2 (Sc) 013 VI 17Car_mul. Cariboptila mulata Botosaneanu, 1977 (Sc) 365 1 031 2,075 Riffle in rainy season 18Ali_ala. Alisotrichia alayoana Botosaneanu, 1977 (Sc, CG) 142 240 19Ali_sp. Alisotrichia sp. 1 (Sc, CG) 3 6 20Ali_sp. y Alisotrichia sp. y (Sc, CG) 27 138 21Leu_sp1. Leucotrichia sp. 1 (Sc, CG) 1 123 VII 22Ato_vin. Atopsyche vinai Sýkora & Botosaneanu, 1973 (Pr) 52 1,316 Riffle in dry season 23Ali_sp2. Alisotrichia sp. 2 (Pc-Hb, Sc, CG) 22 2 24Chi_pul. Chimarra pulchra (Hagen, 1861) (Ft) 131 0 25Chi_gua. Chimarra guapa Botosaneanu, 1977 (Ft) 10 0 26Smi_com. Smicridea comma Banks, 1924 (Ft) 267 170 27Smi_sp1. Smicridea sp. 1 (Ft) 35 2 28Hyd_cub. Hydropsyche cubana (Flint, 1962) (Ft) 454 218 340 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba leaves and bank vegetation were used less by these two species. During the dry season, M. scudderi, a shredder (Ramírez and Gutiérrez Fonseca 2014), primarily inhabited sand microhabitats and a smaller number of riffles. Meanwhile, Helicopsyche sp. primarily used the pool stone, with fewer specimens also inhabiting the sand and riffle microhabitats. Group III was the second most diverse group (Fig. 4C), comprising six species: Hydropsyche sp. 1, Oxyethira sp. 1, Ochrotrichia sp. 1, Xiphocentron cubanum (Botosaneanu, 1993), Antillopsyche sp. 1, and Helicopsyche cf. hageni (Helicopsychidae). The species in this group appear to be stone specialists, occurring both in pools and in riffles. The group also displays a diversity of FFGs, including collector-gatherers, piercers, predators, and filterers. The highest abundance was recorded during the dry season (151 individuals), mainly in the cobble microhabitat of pools. In contrast, only four species were found in the riffle microhabitat during the rainy season, with just a few individuals (18). The most abundant species were Oxyethira sp. (81 individuals) and X. cubanum (46) in the pool cobble microhabitats. Group (IV) was the third most abundant (Fig. 4D), comprising Cernotina sp. 1, Phylloicus chalybeus, Nectopsyche cubana, and Helicopsyche hageni. These species belong to the families Polycentropodidae, Calamoceratidae, Leptoceridae, and Helicopsychidae, respectively. This group showed generalist behavior in microhabitat use and each species belongs to a different functional feeding group. Seasonality did not significantly affect their abundance as values were similar in both the dry and rainy seasons. The only notable exception was Nectopsyche cubana, whose abundance decreased by nearly 50% from the dry to the rainy season. Group (V) (Fig. 4E) contained Cariboptila sp. 1 and Helicopsyche sp. 2 from the families Glossosomatidae and Helicopsychidae, respectively. These morphospecies are both grouped as scrappers FFG, and their preferred microhabitat were pool stones. Helicopsyche sp. 2 was only present in the rainy season. During the rainy season, Cariboptila sp. 1 mainly used the microhabitats of pool stones and riffles, and in the dry season, it also used sand but with fewer specimens. Group (VI) had the highest abundance among all the groups in the k-means analysis (Fig. 4F), formed by Cariboptila mulata, Alisotrichia alayoana, Alisotrichia sp. 1, Alisotrichia sp. y, and Leucotrichia sp. 1. They are members of the Glossosomatidae and Hydroptilidae. The riffles were the preferred microhabitats of this group species. Meanwhile, all members of the group were scrapers. Moreover, in the rainy season, abundance values were twice as high compared to the dry season. Cariboptila mulata was the most abundant species, representing 67% of the group. Seven species were present in Group (VII) (Fig. 4G), the most diverse in our analysis. The species included Atopsyche vinai, Alisotrichia sp. 2, Chimarra pulchra, C. guapa, Smicridea comma, Smicridea sp. 1, and Hydropsyche cubana. They are members of the Hydrobiosidae, Philopotamidae and Hydropsychidae. During the dry season, they preferred almost exclusively the riffle microhabitat. Filterers dominated the functional feeding groups, but predators and scrapers were also present. In the dry season, abundance was higher by 70% over abundance in the rainy season, with H. cubana and S. comma being the most abundant species in both seasons (Table 2) within the group. 341 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba Rainy Season Dry Season A B C D E Figure 4. Percentage of individuals of each species (classified according to k-means groups) found in each microhabitat and season. 348 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba Dudgeon D (1997) Life histories, secondary production and microdistribution of hydropsychid caddisflies (Trichoptera) in a tropical forest stream. Journal of Zoology (London, England) 243(1): 191–210. https://doi.org/10.1111/j.1469-7998.1997. tb05763.x Extence CA (1981) The effect of drought on benthic invertebrate communities in a lowland river. Hydrobiologia 83(2): 217–224. https://doi.org/10.1007/BF00008269 Fisher SG (1983) Succession in streams. In: Barnes JR, Minshall GW (Eds) Stream Ecology: Application and Testing of General Ecological Theory. Plenum Press, New York, 7–27. https://doi.org/10.1007/978-1-4613-3775-1_2 Flint Jr OS (1996) Checklist of the Trichoptera, caddisflies, of Cuba. Cocuyo 5: 15–17. Froehlich CG, Oliveira LG (1997) Ephemeroptera and Plecoptera nymphs from riffles in low-order streamis in southeastern Brazil. In: Landolt P, Sartori M (Eds) Ephemeroptera & Plecoptera: Biology–Ecology–Systematics. MTL, Fribourg, 180–185. Hagen H (1861) Synopsis of the Neurptera of North America. Smithsonian Miscellaneous Collections 4: 347. Henriques Oliveira AL, Nessimian JL (2010) Aquatic macroinvertebrate diversity and composition in streams along an altitudinal gradient in Southeastern Brazil. Biota Neotropica 10(3): 115–128. https://doi.org/10.1590/S1676-06032010000300012 Hernández A, Pérez PJ, Bosh D, Rivero L (1999) Nueva versión de clasificación genética de los suelos de Cuba. Instituto de Suelos, La Habana, Cuba. Holzenthal RW, Thomson RE, Ríos Touma B (2015) Order Trichoptera. In: Thorp JH, Rogers DC (Eds) Thorp and Covich’s Freshwater Invertebrates. Academic Press, San Diego, 965–1002. https://doi.org/10.1016/B978-0-12-385026-3.00038-3 Huryn AD (2009) Aquatic Insects Ecology, Feeding, and Life History. University of Alabama, Tuscaloosa, 132–143. https://doi.org/10.1016/B978-012370626-3.00159-9 INRH (2011) Delegación Provincial de Recursos Hidráulicos Granma. Reporte 2010. Jin HS, Ward GM (2007) Life history and secondary production of Glossosoma nigrior Banks (Trichoptera: Glossosomatidae) in two Alabama streams with different geology. Hydrobiologia 575(1): 245–258. https://doi.org/10.1007/s10750-006-0370-2 Kingsolver JM (1964) New species of Trichoptera from Cuba. Proceedings of the Entomological Society of Washington 66(4): 257–257. Kumanski KP (1987) On caddisflies (Trichoptera) of Cuba. Acta Zoologica Bulgarica 34: 3–35. Lake PS (2003) Ecological effects of perturbation by drought in flowing waters. Freshwater Biology 48(7): 1161–1172. https://doi.org/10.1046/j.1365-2427.2003.01086.x Legendre P, Legendre L (2012) Numerical Ecology (3rd edn). Elsevier Science BV, Amsterdam, 1006 pp. López Del Castillo P, Naranjo López JC, González Lazo DD, Trapero A, Fernández JL, Pérez J (2004) Insectos acuáticos del Parque Nacional “La Bayamesa”, Cuba. Boletin de la SEA 35: 225–231. López Del Castillo P, Naranjo López JC, Fernández Triana JL, González Lazo DD (2007) Caddisflies (Insecta: Trichoptera) in two rivers of eastern Cuba. In: Proceedings of the 12th International Symposium on Trichoptera. The Caddis Press, Columbus, Ohio, 169–173. López Del Castillo P, Gómez Luna LM, López Iborra GM (2024) Microhabitat use and seasonality of mayflies (Ephemeroptera) in two streams in eastern Cuba. Aquatic Insects 45(3): 384–401. https://doi.org/10.1080/01650424.2023.2299810 Martini J, Waringer J (2021) Dynamic microhabitat shifts in space and time of caddisfly larvae (Insecta: Trichoptera) in a first‐order calcareous mountain stream. Biologia 76(9): 2527–2541. https://doi.org/10.1007/s11756-021-00741-w 349 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba Merritt RW, Cummins KW [Eds] (1996) An Introduction to the Aquatic Insects of North America. Kendall Hunt, Dubuque, 862 pp. Minshall GW (1984) Aquatic insect–substratum relationships. In: Resh VH, Rosenberg DM (Eds) The Ecology of Aquatic Insects. Praeger Scientific, New York, 358–400. Morse JC, Frandsen PB, Wolfram G, Thomas JA (2019) Diversity and ecosystem services of Trichoptera. Insects 10(5): 125. https://doi.org/10.3390/insects10050125 Naranjo López JC, González Lazo DD (2005) Orden Trichoptera en Cuba. Boletin de la SEA 1(36): 147–152. Núñez A, Viña Bayes N, Graña A (1989) Regiones naturales antrópicas. In: Hernández JR, Sánchez Herrero A, Propín E, Buznego E, Lorenzo AC, Mon M, Azcue A, et al. (Eds) Nuevo Atlas Nacional de Cuba. Instituto de Geografía, La Habana & Madrid, 232 pp. Oksanen J, Blanchet F, Kindt R, Legendre P, O’Hara R, Simpson G, et al. (2011) Vegan: community ecology package (version 1.17-11). https://cran.r-project.org/web/packages/vegan/index.html Oliveira ALHD, Nessimian JL (2010) Spatial distribution and functional feeding groups of aquatic insect communities in Serra da Bocaina streams, southeastern Brazil. Acta Limnologica Brasiliensia 22(4): 424–441. https://doi.org/10.4322/actalb.2011.007 Ortiz BP, Rivero A (2004) Índices climáticos para la determinación y simulación de las señales de la variabilidad climática en diferentes escalas espacio temporales. Revista Cubana de Meteorología 11(1): 65–75. Poff NL, Ward JV (1989) Implications of streamflow variability and predictability for lotic community structure: A regional analysis of streamflow patterns. Canadian Journal of Fisheries and Aquatic Sciences 46(10): 1805–1818. https://doi.org/10.1139/f89-228 Poole WC, Stewart KW (1976) The vertical distribution of macrobenthos within the substratum of the Brazos River, Texas. Hydrobiologia 50(2): 151–160. https://doi. org/10.1007/BF00019818 Puentes G (2001) Regionalización climática de los macizos montañosos orientales. In: Viña N, Fong A, Maceira D (Eds) Diversidad Biológica de los Macizos Montañosos Orientales. Centro Oriental de Ecosistemas y Biodiversidad (BIOECO), Santiago de Cuba, 14–25. R Core Team (2020) R: a language and environment for statistical computing. RFS Computing, Vienna, Austria. Ramírez A, Gutiérrez-Fonseca PE (2014) Functional feeding groups of aquatic insect families in Latin America: A critical analysis and review of existing literature. Revista de Biología Tropical 62(2): 155–167. https://doi.org/10.15517/rbt.v62i0.15785 Reyes Torres LJ, Ramírez A (2018) Life history and phenology of Phylloicus pulchrus (Trichoptera: Calamoceratidae) in a tropical rainforest stream of Puerto Rico. Revista de Biología Tropical 66(2): 814–825. https://doi.org/10.15517/rbt.v66i2.33411 Reynaga MC (2009) Hábitos alimentarios de larvas de Trichoptera (Insecta) de una cuenca subtropical. Ecología Austral 19(3): 207–214. Reynaga MC, Rueda Martín PA (2014) Trophic analysis of three species of Marilia (Trichoptera: Odontoceridae) from the neotropics. Revista de Biología Tropical 62(2): 543–550. https://doi.org/10.15517/rbt.v62i2.9959 Saito R, Tojo K (2016) Comparing spatial patterns of population density, biomass, and genetic diversity patterns of the habitat generalist mayfly Isonychia japonica Ulmer (Ephemeroptera: Isonychiidae) in the Chikuma–Shinano river basin. Freshwater Science 35(2): 724–737. https://doi.org/10.1086/686537 Springer M (2010) Macroinvertebrados de agua dulce de Costa Rica I, capítulo 7: Trichoptera. Revista de Biología Tropical 58(4): 151–181. 350 ZooKeys 1263: 333–350 (2025), DOI: 10.3897/zookeys.1263.150346 Pedro López Del Castillo et al.: Caddisflies micro-habitat use Cuba Statzner B, rigoux SM, Leichtfried M (2005) Mineral grains in caddisfly pupal cases and streambed sediments: Resource use and its limitation through conflicting resource requirements. Limnology and Oceanography 50(2): 713–721. https://doi. org/10.4319/lo.2005.50.2.0713 Strahler AN (1957) Quantitative analysis of watershed geomorphology. Eos, Transactions American Geophysical Union 38(6): 913–920. https://doi.org/10.1029/ TR038i006p00913 Tomanova S, Goitia E, Helesic J (2006) Trophic levels and functional feeding groups of macroinvertebrates in Neotropical streams. Hydrobiologia 556(1): 251–264. https:// doi.org/10.1007/s10750-005-1255-5 Urbanič G, Toman MJ, Krušnik C (2005) Microhabitat type selection of caddisfly larvae (Insecta: Trichoptera) in a shallow lowland stream. Hydrobiologia 541(1): 1–12. https://doi.org/10.1007/s10750-004-4314-4 Van den Brink FWB, Van der Velde G, Wijnhoven S (2013) Seasonal changes in caddis larvae assemblages in river-floodplain habitats along a hydrological connectivity gradient. Hydrobiologia 716(1): 75–85. https://doi.org/10.1007/s10750-013-1545-2 Vannote RL, Minshall GW, Cummins KW, Sedell JR, Cushing CE (1980) The river continuum concept. Canadian Journal of Fisheries and Aquatic Sciences 37(1): 130–137. https://doi.org/10.1139/f80-017 Vilenica M, Brigić A, Sartori M, Mihaljević Z (2018) Microhabitat selection and distribution of functional feeding groups of mayfly larvae (Ephemeroptera) in lotic karst Habitats. Knowledge and Management of Aquatic Ecosystems 419(17): 1–12. https:// doi.org/10.1051/kmae/2018011 Viña Bayés N (2001) Caracterización geográfica. In: Fong A, Viña N, Maceira D (Eds) Diversidad Biológica de los Macizos Montañosos Orientales. Centro Oriental de Ecosistemas y Biodiversidad (BIOECO), Santiago de Cuba, 4–12. Ward JV (1992) Aquatic Insect Ecology. 1. Biology and Habitat. John Wiley y Sons, _New York 438 pp. Wiggins GB (1996) Larvae of the North American Caddisfly Genera (Trichoptera), 2nd edn. University of Toronto Press, Toronto, 457 pp. https://doi.org/10.3138/9781442623606 Wiggins GB (2007) Caddisflies: Architects under water. American Entomologist 53(2): 78–85. https://doi.org/10.1093/ae/53.2.78