The Smicridea (Smicridea) fasciatella species group (Trichoptera: Hydropsychidae) in Brazil: six new species and new distributional records

The Smicridea (Smicridea) fasciatella species group occurs from the southwestern USA, throughout Central America, the Greater Antilles islands, and most of South America, except for the Chilean subregion. It is characterized by the phallic apparatus being a simple tube with eversible internal sclerites at the apex. The fasciatella group is composed of 61 species, of which only 11 occur in Brazil, mainly in the Atlantic Forest biome in the southeastern region. In order to reduce the Linnean and Wallacean shortfalls for the Smicridea Brazilian fauna, we diagnose, describe, and illustrate males of six new species in the fasciatella group: Smicridea (Smicridea) blahniki Desiderio, Pes & Hamada sp. nov., S. (Smicridea) brevitruncata Desiderio, Pes & Hamada sp. nov., S. (Smicridea) caaguara Desiderio, Pes & Hamada sp. nov., S. (Smicridea) ipiranga Desiderio, Pes & Hamada sp. nov., S. (Smicridea) jeaneae Desiderio, Pes & Hamada sp. nov., and S. (Smicridea) polyacantha Desiderio, Pes & Hamada sp. nov. Additionally, we provide distributional data for S. (Smicridea) albosignata Ulmer, 1907, S. (Smicridea) bivittata (Hagen, 1861), S. (Smicridea) erecta Flint, 1974, S. (Smicridea) obliqua Flint, 1974, S. (Smicridea) paranensis Flint, 1983, and S. (Smicridea) sattleri Denning & Sykora, 1968. The number of S. (Smicridea) species in Brazil increases from 21 to 27 and Smicridea is recorded from the states of Acre, Amapá, and Sergipe for the fi rst time.


Specimen collection, preparation, and observation
Adults were collected with Pennsylvania traps (Frost 1957), UV light pan traps (Calor & Mariano 2012), a white sheet with an LED light, and Malaise traps (Gressitt & Gressitt 1962), or manually with an entomological net. Most of the collected specimens were preserved in 80% or 96% ethanol, while the specimens collected on the white sheets were pinned and preserved dried.
To observe male genital structures, the abdomen of each specimen was removed and cleared using hot 10% KOH as detailed by ; sometimes 85% lactic acid was also used following procedures detailed by Blahnik et al. (2007) in order to evert the endothecal spines of the phallus. After clearing, the abdomen was mounted on a temporary slide using glycerin and examined under a Leica DM5500 B compound microscope. After examination, it was stored in a microvial fi lled with glycerin, together with the remains of the respective specimen in a plastic vial with 80% ethanol.

Illustrations and maps
Photographs of the habitus, head, and forewings were obtained with a Leica DMC4500 digital video camera attached to a Leica M205A stereo microscope or a Leica DFC420 digital video camera attached to a Leica M165C stereo microscope, both using an LED illumination dome (Kawada & Buffi ngton 2016). Stacks of images of each structure were produced at diff erent focal distances; these were then automatically combined into a single image with a greater depth of fi eld using the Helicon Focus ® stacking software (ver. 7.6.4 Pro).
Male genitalia were photographed using a Leica DFC295 video camera attached to a Leica DM5500 B compound microscope, and the stacked images were used as templates to create vector graphic illustrations in Adobe Illustrator ® . Photographs were assembled into plates using Adobe Photoshop ® .
The distribution maps were prepared using QGIS Las Palmas ver. 2.18.10 software (QGIS Development Team 2016). Vector and raster maps used IBGE (2019) and Natural Earth (2020) data. Distribution data for species of Smicridea were compiled from the literature, analyzed specimens, and collection datasets included in the Global Biodiversity Information Facility (GBIF 2020).

Morphological terminology, descriptions, and material examined
Morphological terminology follows mostly Blahnik (1995) for the male genitalia and Wells & Neboiss (2018) for the head setal warts.
Species descriptions were made using the DELTA (Description Language for Taxonomy) software (Dallwitz et al. 1999) to promote consistency and standardization in descriptive taxonomy. The style of descriptions follows Desiderio et al. (2020).
Lists of material examined were prepared using the AUTOMATEX macro in Microsoft Excel (Brown 2013).

Repositories
Types and other material examined are deposited in the following collections: Diagnosis Smicridea blahniki sp. nov. is similar to S. erecta and S. jeaneae Desiderio, Pes & Hamada sp. nov. based on the strong, pointed apicomesal lobes on tergum X. Of these species, the new species is most similar to S. erecta, because the apicomesal lobes in both species touch distally and each lobe bears one pair of short, thin apical setae in dorsal view, while in S. jeaneae sp. nov., the apicomesal lobes do not touch distally and each has three medium-sized, thin apical setae. Smicridea blahniki sp. nov. has one pair of long, stout subapical setae on the apicomesal lobe of tergum X, whereas S. erecta has one pair of mediumsized, thin setae (absent in S. jeaneae sp. nov.). The basal segment of the inferior appendage in the new species has fi ve stout apical spine-like setae, one long seta on the dorsal surface and four medium-sized on the mesal margin, whereas S. erecta and S. jeaneae sp. nov. have three stout apical spine-like setae (one dorsal and two on the internal margin). Additionally, the apical segment of the inferior appendage in S. blahniki sp. nov. is thick, slightly curved medially, and has a rounded apex, while in S. erecta and S. jeaneae sp. nov., this segment is slender, strongly curved medially, and has a slightly acute apex. Furthermore, S. blahniki sp. nov. can be recognized by the stethoscope-shaped phallotremal sclerite of the phallic apparatus in dorsal view, while in S. erecta this sclerite is ribbon-shaped and in S. jeaneae sp. nov. it is U-shaped in the proximal region and hastate in the distal region.

Etymology
The specifi c epithet is in honor of Dr Roger J. Blahnik (University of Minnesota, USA) in recognition of his contributions to the taxonomy and systematics of Neotropical caddisfl ies. C . General color brown (in alcohol) (Fig. 1A). Antennae yellowish brown (Fig. 1A). Head brown, denuded ; dorsally with weakly-pronounced median suture; with fi ve setal warts; anteromesal wart small, rounded; anterolateral pair undivided, weakly delimited; posterolateral pair large, ovoid (Fig. 1C). Maxillary palp with segments 1-4 increasing gradually about 0.8 × in size toward more apical segments, segment 5 long, about 1.2 × as long as all previous segments combined (Fig. 1B). Thorax pale brown, without setae (old specimens); legs brown, with mid leg tarsi yellowish (Fig. 1A). Wing venation typical for subgenus . Forewings, in alcohol, uniformly brown, with small white spot on apex of Cu2 (Fig. 1D). Sternum V with anterolateral glandular processes as long as sternum. Abdominal segments VI and VII with 2 pairs of internal glands, both pairs longer than their containing segments (Fig. 1F). M (Fig. 2). Segment IX in lateral view with anterolateral margin nearly straight, strongly sclerotized (Fig. 2A); posterodorsal margin bearing small spicules, distributed in a subtriangular band (Fig. 2B). Tergum X elongate; in lateral view, ventrolateral margin strongly sclerotized, with four mediumsized, thin setae on apical third; apex with sharp notch, ending in slender, straight lobe ( Fig. 2A); in dorsal view, lateral margin rounded; dorsomesal setose area bearing three medium-sized, thin setae; divided apicomesally by V-shaped incision about ¼ its length; apex of tergite rounded, with strong, pointed apicomesal lobe, touching distally, each bearing one pair of long, stout subapical setae and one pair of short, thin apical setae (Fig. 2B). Inferior appendages 2-segmented; basal segment long, slightly infl ated distally, covered with long, thin setae, with a group of stout apical spine-like setae, one long seta dorsally and four medium-sized setae on mesal margin; apical segment short, about ⅓ as long as basal segment, thick, slightly curved medially, apex rounded with numerous short, thin setae on mesal margin (Fig. 2B). Phallic apparatus long and tubular; basal section, in lateral view, enlarged, 4 × diameter of phallotheca at its narrowest point, forming an angle of about 93° with apical section; median section of phallotheca straight, with dorsal periphalic cap sclerotized, slender, and upturned (Fig. 2C); apex enlarged, without processes or plates; endothecal membranes without spines; phallotremal sclerite, in dorsal view, distinct, strongly sclerotized, stethoscope-shaped, proximal region slightly bifi d and distal region with acute arms

Bionomics
The males of S. blahniki sp. nov. were collected near a 1 st order stream, where the predominant vegetation is highland savanna with a minimum altitude of 1112 m a.s.l. (Fig 4A). This stream is located in the Serra do Aracá mountain range, northern Brazilian Amazon. The water in the stream was black and acidic (pH 4.6), with low conductivity (10 μScm -1 ), and a water temperature of 22°C. The stream was about 0.5 m wide, 0.30 m deep, with rocky bottoms characterized by boulders and slow-fl owing water. Diagnosis Smicridea brevitruncata sp. nov. belongs to the albosignata complex of Flint (1981). It is most similar to S. riita Flint, 1981 from Venezuela based on the obliquely truncated apex of tergite X in dorsal view. However, in S. brevitruncata sp. nov. the apical segment of the inferior appendage is thick and short, and the truncated apex has numerous medium-sized, acuminate setae on the mesal margin, whereas in S. riita it is long, slender, with an acute apex, bearing numerous short, thin setae. Furthermore, the new species can be diagnosed by the basal segment of the inferior appendage having four long, stout apical spine-like setae, two dorsal and two on the mesal margin, and by the shape of the phallotremal sclerite, mainly in dorsal view. Additionally, S. brevitruncata sp. nov. diff ers from all other species of S. (Smicridea) by the absence of the internal reticulate sacs on abdominal segments VI and VII.

Etymology
The specifi c epithet is an allusion to the characteristics of the inferior appendages, which are short and apically truncated. Derived from the Latin words, ʻbreviʼ = ʻshortʼ and ʻtruncataʼ = ʻpiece cut off , tip, endʼ. C . General color dark brown (in alcohol) (Fig. 5A). Antennae brown (Fig. 5A). Head dark, with grayish to black setae on dorsum ; dorsally with slightly pronounced median suture; with fi ve setal warts; anteromesal wart small, rounded; anterolateral pair undivided, weakly delimited; posterolateral pair large, ovoid (Fig. 5C). Maxillary palp with segments 1-4 increasing gradually about 0.8 × in size  toward more apical segments, segment 5 long, about 1.2 × as long as all previous segments combined (Fig. 5B). Thorax dark brown, covered with brown setae; legs brown, with mid-leg tarsi yellowish (Fig. 5A). Wing venation typical for subgenus . Forewings, in alcohol, uniformly fuscous, faintly marked with transverse band subapically on pterostigma reaching base of R4+5 and small spot on apex of Cu2 (Fig. 5D). Sternum V with anterolateral glandular processes slightly shorter than sternum. Abdominal segments VI and VII without pairs of internal reticulate sacs (Fig. 5F). Tergum X elongate; in lateral view, ventrolateral margin strongly sclerotized, with 10 to 12 short, stout setae on apical third; apex produced and upturned (Fig. 6A); in dorsal view, lateral margin slightly rounded; dorsomesal setose area bearing 10 medium-sized, thin setae; divided apicomesally by V-shaped incision about ⅓ its length; apex of tergite obliquely truncated, with about three short, thin setae (Fig. 6B). Inferior appendages 2-segmented; basal segment long, slightly infl ated distally, covered with long, thin setae, with four long, stout apical spine-like setae, two dorsal and two on mesal margin; apical segment short, about ¼ as long as basal segment, thick, slightly curved medially, apex truncated with numerous medium-sized, acuminate setae on mesal margin (Fig. 6B). Phallic apparatus long and tubular; basal section, in lateral view, enlarged, 4 × diameter of phallotheca at its narrowest point, forming an angle of about 113° with apical section; median section of phallotheca straight, without processes (Fig. 6C); apex enlarged, with pair of claw-shaped internal dorsolateral plates, longer than basal width, curved mesally and directed posterad; endothecal membranes without spines; phallotremal sclerite, in dorsal view, distinct, strongly sclerotized, proximal region with bifi d base and pointed apex, distally with pair of subtriangular, toothlike structures, which are overlapped by pair of slender, curved mesad processes ; in lateral view, proximal region with small middorsal lobe, distally directed downward (Fig. 6C). Ejaculatory duct of endophallus, in lateral view, distinct, slightly sclerotized (Fig. 6C).

Bionomics
This species was collected near a large fast-fl owing river about 20 m wide, located at 1022 m a.s.l. in the southern part of the Brazilian Atlantic Forest (Fig 4B). In this region the predominant vegetation is Mixed Ombrophilous Forest composed mainly of Araucaria angustifolia (Bertol.) Kuntze (Araucariaceae), with intense agricultural and pasture activities.

Diagnosis
This new species is another member of the albosignata complex; it is most similar to the nominal species S. albosignata based on the enlarged, slightly bifi d apex of the apical segment of the inferior appendage. However, the lateral margin of tergum X in the new species is rounded in dorsal view, while in S. albosignata this structure is straight. Both species also diff er in the shape of the tergum X apex: in S. albosignata it is acute in dorsal view, while in S. caaguara sp. nov. it is rounded. Additionally, S. albosignata has the apex of tergum X with a subtriangular apicomesal fl ap directed laterad in dorsal view, while in S. caaguara sp. nov. this fl ap is absent. The new species also can be recognized by the distinct and complex phallotremal sclerite in dorsal view.

Etymology
The specifi c epithet is a reference to the indigenous people known as caaguaras, fi rst inhabitants of the São Francisco de Paula municipality, which is the type locality of this species. This name is used as a noun in apposition. Thorax dark brown, covered with brown setae; legs brown, with mid-leg tarsi yellowish (Fig. 7A). Wing venation typical for subgenus . Forewings, in alcohol, grayish to dark brown, with white transverse band subapically on pterostigma reaching base of R4+5 and small white spot on apex of Cu2 (Fig. 7D). Sternum V with anterolateral glandular processes slightly longer than sternum. Abdominal segments VI and VII with 2 pairs of internal glands, both pairs longer than their segments (Fig. 7F). M (Fig. 8). Segment IX in lateral view with anterolateral margin nearly straight, weakly sclerotized (Fig. 8A); posterodorsal margin bearing small spicules, distributed in two dark, narrow parallel bands (Fig. 8B). Tergum X elongate; in lateral view, ventrolateral margin strongly sclerotized, with eight to 10 medium-sized, thin setae on apical third; apex produced and upturned (Fig. 8A); in dorsal view, lateral margin rounded; dorsomesal setose area bearing six to eight medium-sized, thin setae; divided apicomesally by V-shaped incision about ⅓ its length; apex of tergite rounded, with about three short, thin setae (Fig. 8B). Inferior appendages 2-segmented; basal segment long, slightly infl ated distally, covered with long, thin setae, with pair of long, stout apical spine-like setae on mesal margin; apical segment short, about ½ as long as basal segment, thick, curved medially, apex enlarged and slightly bifi d with numerous short, acuminate setae on mesal margin (Fig. 8B). Phallic apparatus long and tubular; basal section, in lateral view, enlarged, 3 × diameter of phallotheca at its narrowest point, forming an angle of about 105° with apical section; median section of phallotheca straight, with dorsal periphalic cap sclerotized and fl attened (Fig. 8C); apex enlarged, with pair of claw-shaped internal dorsolateral plates, longer than basal width, curved mesally and directed posterad; endothecal membranes without spines; phallotremal sclerite, in dorsal view, distinct, strongly sclerotized, proximal region slender, rod-like, divided distally in two subtriangular lateral fl aps ; in lateral view, proximal region with small middorsal lobe, distally directed downward (Fig. 8C). Ejaculatory duct of endophallus, in lateral view, distinct, slightly sclerotized (Fig. 8C).

Bionomics
The males of S. caaguara sp. nov. were collected near medium-sized streams, about 5.0 m wide, located in the Pampa-Atlantic Forest transition zone in the state of Rio Grande do Sul. The vegetation of the collections sites varies from Grassy-Woody Steppe (Arroio Lageado -Rio dos Sinos Basin, Fig 4C) to Seasonal Deciduous Forest (Arroio Teotônia -Rio Jacuí Basin, Fig 4D). The streams have bedrock and fast-fl owing water, with cloudy and alkaline water (pH 7.9), conductivity 25-50.4 μScm -1 , and water temperatures of 15-16.4°C. Distribution (Fig. 3) Brazil: Atlantic Forest (Rio Grande do Sul).

Diagnosis
This new species can be assigned to the fasciatella complex. Smicridea ipiranga sp. nov. is similar to S. varia (Banks, 1913), S. lacanha Hamilton, 1986, andS. catherinae Blahnik, 1995 based on the shape of the apical segment of the inferior appendage, which is slender, strongly curved medially and has a pointed apex. These four species also have a single long, stout apical spine-like seta on the internal margin of the basal segment of the inferior appendage. However, S. ipiranga sp. nov. can be distinguished from its congeners by its diminutive size (on average 3.55 mm); the most similar species in the complex are twice its size on average: S. catherinae (about 6.0 mm), S. lacanha (about 7.9 mm), and S. varia (about 6.15 mm). Additionally, the new species has the phallotremal sclerite with the distal region M-shaped in dorsal view, while in S. catherinae, S. lacanha, and S. varia, this sclerite is Y-shaped.

Etymology
The specifi c epithet is a reference to the Ipiranga Stream, located in the Reserva Florestal Adolpho Ducke, where the type specimen was collected. This name is used in apposition. C . General color dark brown (in alcohol) or black (pinned) (Fig. 9A). Antennae yellowish brown (Fig. 9A). Head dark, with white setae on dorsum (Fig. 9B-C); dorsally with weakly-pronounced median suture; with fi ve setal warts; anteromesal wart small, rounded; anterolateral pair undivided, weakly delimited; posterolateral pair large, ovoid (Fig. 9C). Maxillary palp with segments 1-4 increasing gradually about 0.8 × in size toward more apical segments, segment 5 long, about 1.2 × as long as all previous segments combined (Fig. 9B). Thorax dark brown, covered with pale brown setae; legs dark brown, with mid-leg tarsi pale yellowish (Fig. 9A). Wing venation typical for subgenus (Fig. 9D-E). Forewings, in alcohol, brown, with white transverse band subapically on pterostigma reaching base of R4+5 and small white spot on apex of Cu2 (Fig. 9D); pinned specimens with 2 transversal bands of white setae, one continuous band in proximal portion reaching from base of subcostal vein to apex of 1A, the other band discontinuous in distal portion reaching from pterostigma to apex of Cu1; apex of wing with fringe of white setae (Fig. 9A). Sternum V with anterolateral glandular processes slightly shorter than sternum. Abdominal segments VI and VII with 2 pairs of internal glands, both pairs longer than their containing segments (Fig. 9F).  (Fig. 10). Segment IX in lateral view with anterolateral margin straight, strongly sclerotized (Fig. 10A); posterodorsal margin bearing small spicules, distributed in two subquadrate bands (Fig. 10B). Tergum X elongate; in lateral view, ventrolateral margin strongly sclerotized, with small subapical lobe; apex produced and upturned (Fig. 10A); in dorsal view, lateral margin slightly sinuous; dorsomesal setose area bearing fi ve medium-sized, thin setae; divided apicomesally by V-shaped incision about ¼ its length; apex of tergite rounded, with large globose lobe and about eight short, stout setae (Fig. 10B). Inferior appendages 2-segmented; basal segment long, slightly infl ated distally, covered with long, thin setae, with one long, stout apical spine-like seta on mesal margin; apical segment short, about ⅓ as long as basal segment, slender, strongly curved medially, apex acute with numerous short, thin setae on mesal margin (Fig. 10B). Phallic apparatus long and tubular; basal section, in lateral view, enlarged, 2 × diameter of phallotheca at its narrowest point, forming an angle of about 116° with apical section; median section of phallotheca straight, without processes (Fig. 10C); apex slightly enlarged, without processes or plates; endothecal membranes without spines; phallotremal sclerite, in dorsal view, distinct, strongly sclerotized, proximal region enlarged and narrowing mesally, distal region M-shaped ; in lateral view, aciculate (Fig. 10C). Ejaculatory duct of endophallus, in lateral view, distinct, slightly sclerotized (Fig. 10C). When endothecal membranes are everted, phallotremal sclerite divided into two disconnected parts, one long, apicomesal, rod-like, and the other with a pair of comma-shaped sclerites, distally slightly curved (Fig. 10F).

Bionomics
The males of S. ipiranga sp. nov. were collected near streams in the central Amazon, mainly in a 2 nd order stream (Igarapé Ipiranga) located in the Reserva Florestal Adolpho Ducke, an area of primary Amazon rainforest of about 100 km 2 that is under the protection of the Instituto Nacional de Pesquisas da Amazônia (INPA). The riparian vegetation along this stream is about 2.4 m wide and 0.29 m deep, with a sandy bottom, and a medium-speed water fl ow (Fig 4E). Its water is clear and acidic (pH 5.1), oxygen solubility levels are high (6.9 mgL), electrical conductivity is low (8.5 μScm -1 ), and the average temperature is 25.6°C. Specimens of S. ipiranga sp. nov. were also collected in medium-sized streams in the municipality of Presidente Figueiredo (Corredeira da Dona Maroca and Igarapé da Onça), 3.5-12.5 m in width, 0.19-0.44 m deep, with fast-fl owing water. These streams were used for recreation (swimming and camping), with rocky bottoms characterized by black and acidic water (pH 4.2-4.8), low electrical conductivity (10.9 μScm -1 ), and a water temperature of 25ºC. Diagnosis Smicridea jeaneae sp. nov. also belongs to the albosignata complex. Among the species in this complex, it is most similar to S. bulbosa Flint, 1974 from Suriname based on the strong, parallel, pointed apicomesal lobes on tergum X and the short, slender apical segment of the inferior appendage with slightly acute apex. In both species the claw-shaped internal dorsolateral plates of the phallic apparatus are as long as the basal width, but in S. jeaneae sp. nov. the plates are straight mesally, while in S. bulbosa they are curved. In addition, the phallotremal sclerite of the phallic apparatus has a hastate shape distally in dorsal view and is kidney-shaped in lateral view in the new species, but is strap-like in dorsal view and rod-like in lateral view in S. bulbosa. Smicridea jeaneae sp. nov. can also be recognized by the three stout apical spine-like setae on the basal segment of the inferior appendage, one long seta dorsally and two mediumsized setae on the mesal margin.

Etymology
This new species is named in honor of Dr Jeane Marcelle Cavalcante do Nascimento (Instituto Nacional de Pesquisas da Amazônia, INPA -Brazil) for her valuable friendship, cooperation in the fi eld and the laboratory, and in gratitude for her help in collecting material during the doctoral studies of the fi rst author.

Bionomics
The only male specimen of S. jeaneae sp. nov. was collected near the Oiapoque River, at a place known as "Buraco do Cristal"; this river is about 400 m in width. The prevalent vegetation is Dense Ombrophilous Lowland Forest. The collection site is located in the northern-most part of the Brazilian Amazon. Distribution (Fig. 3) Brazil: Amazon (Amapá).

Diagnosis
This new species is a distinct member of the fasciatella group due the presence of small endothecal spines on the apex of the phallic apparatus. Smicridea polyacantha sp. nov. and S. paranensis are the sole species in the group that share this character. However, the new species has 40 conical endothecal spines grouped in a V-shaped arrangement in dorsal view, while S. paranensis has numerous fi ne spines organized in four rows. Additionally, the apex of the apical segment of the inferior appendage is rounded in the new species and obliquely truncated in S. paranensis. The phallotremal sclerite has a proximal region that is bifi d, curved downward in lateral view, and a dorsolateral projection on the distal region that is unique in S. polyacantha sp. nov.

Etymology
The specifi c epithet is an allusion to the large number of endothecal spines present on the apex of the phallus. Derived from the Greek words, ʻpolyʼ = ʻveryʼ or ʻmanyʼ and ʻakanthaʼ = ʻspineʼ. C . General color dark brown (in alcohol) (Fig. 13A). Antennae yellowish brown (Fig. 13A). Head dark brown, with brown setae on dorsum (Fig. 13B-C); dorsally with weakly pronounced median suture; with fi ve setal warts; anteromesal wart small, rounded; anterolateral pair undivided, weakly delimited; posterolateral pair large, ovoid (Fig. 13C). Maxillary palp with segments 1-4 increasing gradually about 0.8 × in size toward more apical segments, segment 5 long, about 1.2 × as long as all previous segments combined (Fig. 13B). Thorax dark brown, covered with pale-brown setae; legs brown, with mid-leg tarsi yellowish (Fig. 13A). Wing venation typical for subgenus (Fig. 13D-E). Forewings, in alcohol, brown, with white transverse band subapically on pterostigma reaching base of R4+5 and small white spot on apex of Cu2 (Fig. 13D). Sternum V with anterolateral glandular processes slightly shorter than sternum. Abdominal segments VI and VII with 2 pairs of internal glands, both pairs as long as their containing segments (Fig. 13F).   (Fig. 14). Segment IX in lateral view with anterolateral margin nearly straight, weakly sclerotized (Fig. 14A); posterodorsal margin bearing small spicules, distributed in two rounded bands (Fig. 14B). Tergum X elongate; in lateral view, ventrolateral margin slightly sclerotized, with fi ve long, thin setae on apical third; apex produced and upturned (Fig. 14A); in dorsal view, lateral margin straight; dorsomesal setose area bearing two medium-sized, thin setae; divided apicomesally by a V-shaped incision about ½ its length; apex of tergite slightly pointed, with small apicomesal lobe and about six mediumsized, thin setae (Fig. 14B). Inferior appendages 2-segmented; basal segment long, slightly infl ated distally, covered with medium-sized, thin setae, with one long, stout subapical spine-like seta on mesal margin; apical segment short, about ⅓ as long as basal segment, thick, slightly curved medially, apex rounded with numerous short, thin setae on mesal margin (Fig. 14B). Phallic apparatus long and tubular; basal section, in lateral view, enlarged, 3 × diameter of phallotheca at its narrowest point, forming an angle of about 116° with apical section; median section of phallotheca straight, with dorsal periphalic cap sclerotized, slender and upturned (Fig. 14C); apex slightly enlarged, without processes or plates; endothecal membranes with 40 small, conical spines, grouped in a V-shaped arrangement and directed posterad; phallotremal sclerite, in dorsal view, distinct, strongly sclerotized, proximal region bifi d, curved downward in lateral view and directed posterad, distal region with dorsolateral projections more than half length of sclerite, broad apically, folding and touching dorsomesally, and with another pair of slender ventrolateral projections curved downward and directed anterad ; in lateral view, proximal and distal regions hooked, not parallel in proximal region (Fig. 14C). Ejaculatory duct of endophallus, in lateral view, indistinct (Fig. 14C). When endothecal membranes are everted, phallotremal sclerite modifi ed into two dorsolateral slender lobes, forked at apex, connected to a dorsomesal ring, and apical spines divided into two lateral lobes with 20 spines each, in lateral view, directed anterad (Fig. 14F-G).

Bionomics
This species was collected near medium-sized streams with slow fl ow in central Amazonia. The streams were strongly impacted by human activities, especially recreational use (swimming) and road building (Fig. 4F). These streams have sandy bottoms and are characterized by Dense Ombrophilous Forest Lowlands to Dense Ombrophilous Forest. Only one male of S. polyacantha sp. nov. was collected near a large river (Rio Papagaio) with fast-fl owing water and a rocky bottom, located in the western part of the Brazilian Cerrado, where the vegetation is composed by Primary Savanna, Seasonal-Forest Savanna, and Wooded Savanna. The surroundings of this region are also strongly impacted by agricultural and pasture activities. Distribution (Fig. 3) Brazil: Amazon (Amazonas), Cerrado (Mato Grosso).

Remarks
This species is widely distributed in Central America and some countries in northern South America (Colombia, Peru, Suriname, Venezuela) (Fig. 15B) (Holzenthal & Calor 2017). In Brazil, S. bivittata has been recorded only from the Cerrado biome in the states of Minas Gerais and São Paulo (Southeast region), and the Amazon biome in the state of Pará (North region) (Santos et al. 2020a). Nevertheless, this species was incorrectly recorded from the state of Pará by Paprocki et al. (2004) in their checklist of Brazilian Trichoptera because Paprocki's work was based on data from the literature and until then no previous study had reported the occurrence of S. bivittata from either the state of Pará or Brazil. This species was recorded in Brazil for the fi rst time by , from the state of Minas Gerais. Thus, this study provides the fi rst record of S. bivittata from the Amazon biome in the state of Pará, based on specimens collected in a small stream located in the Floresta Nacional do Tapajós, a protected Amazon forest area.
The range of S. bivittata is extended in the Cerrado to the Distrito Federal and the state of Tocantins, and in the Amazon biome to the state of Amazonas. In addition, this study provides the fi rst records of this species in the Atlantic Forest biome in the states of Alagoas, Bahia, Espírito Santo, Paraíba, and Pernambuco (Fig. 15B). Flint, 1974 Fig. 16A Smicridea erecta Flint, 1974b: 87 (type locality: Suriname, Nassau Mountains, Km 11.3, creek;RNH;♂).

Discussion
Although Smicridea is highly diverse and widely distributed in Brazil, its geographic distribution is poorly known, and its diversity may be underestimated because some Brazilian states, regions, and biomes have not been sampled yet. Some recent contributions have been made to reduce the biodiversity shortfalls regarding the Brazilian Smicridea fauna (Rocha et al. 2016;Alves et al. 2018;Vilarino et al. 2019;Desiderio et al. 2020Desiderio et al. , 2021. In this paper, we describe six new species of S. (Smicridea), signifi cantly increasing the number of species of this subgenus in Brazil from 21 to 27. We also provide the fi rst records of the genus Smicridea for the Brazilian states of Acre, Amapá, and Sergipe; and of S. (Smicridea) for the states of Rio Grande do Sul and Santa Catarina. The records of S. jeaneae sp. nov. in Amapá State and S. caaguara sp. nov. in Rio Grande do Sul State are the northernmost and southernmost records of the genus in Brazil, respectively.
Genital structures have been remarkably useful for diff erentiating species in the S. fasciatella group treated here. The number, type, and position of the setae and the shape of the apex of tergum X are the most useful characters. Additionally, the number of stout apical spine-like setae on the basal segment of the inferior appendage and the shape of the apical segment, which has acuminate setae or thin setae on its apex, are useful characters to discriminate species in this group. The shape of the phallotremal sclerite of the phallus (especially as viewed dorsally), used by Flint (1974a) to diff erentiate species in S. fasciatella group from North and Central America, also proved to be a useful character for diagnosing Brazilian species in this group. Even though the shape of the phallotremal sclerite has not been useful for separating species in the S. fasciatella complex (Blahnik 1995), we believe that this is one of the most important diagnostic characters for identifying species in the fasciatella species group as a whole.
In this study, some new species do not present diagnostic characters that would allow us to place them in one of the two currently recognized species subgroups (albosignata and fasciatella). However, we note that they share characters that are unusual within the S. fasciatella group, such as a distinct apicomesal lobe on each tergite X in S. blahniki sp. nov., S. erecta, and S. jeaneae sp. nov., and the presence of endothecal spines in S. paranensis and S. polyacantha sp. nov. These shared characteristics may be an indication of the existence of a new subgroup within the S. fasciatella group.
Curiously, the reticulate sacs are absent in S. brevitruncata sp. nov. This fact is unique within the subgenus Smicridea. In S. patinae Pauls, Holzenthal, 2010 andS. microsaccata Flint, 1991 from Chile and Colombia, respectively, only a pair of reduced internal glands are present in segment VII. The internal glands in abdominal segments VI and VII seem to be a plesiomorphic character in S. (Smicridea), as they also occur in the closely related Smicrideinae genus Smicrophylax Neboiss, 1977, and in the Diplectroninae genera Austropsyche Banks, 1939, Arcyphysa Wells & Neboiss, 2018, and Diplectrona Westwood, 1839(Neboiss 1977Wells & Neboiss 2018).
Because S. (Smicridea) groups or subgroups have never been treated under a cladistic approach, a phylogenetic study would be very relevant to evaluate how these characters are related to each other. This is needed to delimit the already-proposed taxonomic subgroups (e.g., albosignata, fasciatella complexes) and to establish any potentially new subgroups. Nevertheless, this is outside of the scope of this paper because we only focused on the description of new species and the distribution of Brazilian species in the S. fasciatella group.
This study highlights the need for more taxonomic studies focused on S. (Smicridea) in Brazil in general, especially in the poorly sampled Caatinga, Pampas, and Pantanal biomes. The S. (Smicridea) species seem to prefer small to medium-sized streams based on the bionomic data for the new species and the collection data on the known species that are treated here. Future collections should therefore be directed to poorly sampled environments. Additionally, there are many undescribed species deposited in diff erent entomological collections worldwide (Santos et al. 2020b). Therefore, more taxonomists are needed to make all of this unknown diversity available.
(UFRJ) for their assistance and the loan of valuable specimens deposited in their collections. We thank Marcos Silveira and Martin Acosta Oliveira (UFAC) for assistance in obtaining collecting licenses and for help in the fi eldwork in the PPBio grid in Acre state. We also thank the Invertebrate Collection of INPA for allowing the use of photographic equipment. Conselho Nacional de Desenvolvimento Científi co e Tecnológico (CNPq) provided a doctoral fellowship for GRD (141875/2018-442 6) and a research fellowship for NH (307849/2014-7; 308970/ 2019-5). AMP received post-doctoral fellowships from PCI/INPA (301401/2019-5) and PDS/CNPq (165223/2017-0). We are grateful to Philip M. Fearnside for revising an earlier version of this manuscript.