Revision of the Neotropical caddisfly genus Itauara Müller, 1888 (Trichoptera, Glossosomatidae)

Abstract Systematics of the genus Itauara Müller, 1888 are reviewed. A generic diagnosis, illustrations, and descriptions are provided for males. The genus can be identified by several features of the male genitalia including an extremely reduced phallobase and a phallic apparatus that consists of a sclerotized dorsal sheath covering a very membranous ventral portion. A total 18 species are described as new: Itauara alexanderi sp. n.(Brazil), Itaura bidentata sp. n. (Guyana), Itaura blahniki sp. n. (Brazil) Itaura charlotta sp. n. (Brazil), Itaura emilia sp. n. (Brazil), Itaura flinti sp. n. (Brazil), Itaura guyanensis sp. n. (Guyana), Itaura jamesii sp. n. (Brazil), Itaura julia sp. n. (Brazil), Itaura lucinda sp. n. (Brazil), Itaura ovis sp. n. (Guyana, Venezuela), Itaura peruensis sp. n. (Peru), Itaura rodmani sp. n. (Brazil), Itaura simplex sp. n. (Brazil), Itaura spiralis sp. n. (Guyana), Itaura stella sp. n. (Brazil), Itaura tusci sp. n. (Brazil), and Itaura unidentata sp. n. (Guyana). These additions bring the total fauna of Itauara to 22 species.


introduction
The genus Itauara Müller, 1888, belongs to the saddle-, or tortoise-case making caddisfly family Glossosomatidae. The name Itauara comes from the Tupi-Guarani language and roughly translates to "born from rock," likely referring to glossosomatid larval cases, often found conspicuously on the surface of submerged rocks. Itauara larvae construct rather loose, easily deformable cases of large and small grains of sand that are vaulted dorsally, and almost flat ventrally (Angrisano 1993). In southeastern Brazil and surrounding regions in Argentina and Uruguay, larvae are known to occur in sandy bottom streams with scarce vegetation where they attach their cases to Characeae algae (Angrisano 1993).
Like other members of Protoptilinae, Itauara adults are minute, usually less than 3 mm in size. Their wings are brownish and may have a conspicuous white spot at the arculus or transverse line along the anastomosis (Fig. 1). Itauara are rather rare in occurrence; indeed, several of the new species described here are known from only a single specimen. The 4 known species of Itauara occur only in South America, with 3 of these [I. brasiliana (Mosely, 1939), I. guarani (Angrisano, 1993), and I. plaumanni (Flint, 1974)] endemic to southeastern Brazil and surrounding regions of Argentina and Uruguay. A fourth species, I. amazonica (Flint, 1971), is known from Amazonas state, Brazil.
A recent survey of the Trichoptera of southeastern Brazil by researchers at the University of Minnesota yielded numerous new species of Itauara and impetus for this study. Dr. Oliver S. Flint, National Museum of Natural History, Smithsonian Institution, generously provided several additional undescribed species from various locales in South America for inclusion in this study. In this paper, we determine the homologies and establish standardized terminology of the male genitalic structures among species. Additionally, we re-describe and illustrate the 4 known Itauara species and describe 18 new species from southeastern Brazil, Peru, Guyana, and Venezuela, bringing the total number of species to 22. These species are here assigned to 2 species groups and listed in Table 1. Finally, we provide a key to the males of Itauara.

Specimen preparation and observation
To observe certain structural features of the male genitalia, soft tissues were cleared using a lactic acid method outlined in detail by Holzenthal and Anderson (2004) and Blahnik et al. (2007). For some specimens, the entire individual was cleared (after removing the wings) to more easily observe external structures obscured by setae, such as thoracic warts. Specimens that were over-cleared or lightly sclerotized were stained. Such specimens were immersed in a small watch-glass containing Chlorazole Black E (Sigma Chemical Co.) dissolved in glycerin for 15 minutes to several hours, depending on the size and condition of the specimen. Stained specimens were then rinsed in distilled water to remove any excess stain. Specimens were examined in a small watch-glass containing glycerin using an Olympus SZX12 dissecting microscope or Olympus BX41 compound microscope. To observe wing venation, wing mounts of each species were prepared following the protocols of Blahnik and Holzenthal (2004). Table 1. Itauara species groups (Trichoptera: Glossosomatidae: Protoptilinae).

Illustrations, descriptions, and identification key
Pencil sketches of the male genitalia were completed using either an optical grid on a dissecting microscope, or camera lucida (drawing tube) mounted on a compound microscope. Pencil sketches were scanned digitally, and then placed as a template layer in Adobe Illustrator® for final rendering. Wing preparations were digitally photographed using a Leica EC3 digital camera mounted on an Olympus SZX12 dissecting microscope. Digital images were then placed as a template layer in Adobe Illustrator® for final rendering. Descriptions of species and generation of the identification key were facilitated by using the software package DELTA (DEscriptive Language for Taxonomy) (Dallwitz 1980;Dallwitz et al. 1993 onwards;1999 onwards). Females, with similar size and coloration as males that were collected at the same time and locality, are listed as paratypes under the material examined for some species. Previous experience has shown that having presumptively associated female specimens may be useful for future associative studies. However, since there is some uncertainty of association, we have deferred descriptions of females.

Depositories
Types and additional material examined for this study are deposited at the British Museum of Natural History, London, UK (BMNH); the National Museum of Natural History, Washington, DC, USA (NMNH), the Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenas Aires, Argentina (MACN), the Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZUSP), and the University of Minnesota Insect Collection, Saint Paul, USA (UMSP). All specimens or lot of alcohol specimens examined in this study were affixed with a barcode label with a unique 9 digit alphanumeric code starting with the prefix UMSP. This prefix indicates that the specimen has been databased at UMSP, but it is not meant to imply possession by UMSP. Specimen-level taxonomic, locality, and other information, are stored in the University of Minnesota Insect Collection Biota Trichoptera Database using the software program Biota (Colwell 2003), and can be accessed at http://www.entomology. umn.edu/museum/databases/BIOTAdatabase.html.

systematics
Until recently, the generic status of Itauara was uncertain. Müller (1888) first used the name Itauara in a discussion of larval morphology, but he did not include any species or illustrations. In a later, posthumous work (Müller 1921), he provided sketches of the female forewing venation and some larval structures. Ulmer (1957) thought that Müller's illustrations resembled those of other genera in Protoptilinae of South American origin, and suggested that the larvae be split into different species and perhaps even different genera. He also noted that Müller's forewing illustration completely matched that of Antoptila brasiliana Mosely, 1939(Ulmer 1957. In his studies of Trichoptera collected from the Amazon, Marlíer (1964) later described some Protoptilinae larvae and female pupae and attributed them to Itauara. However, since the pupae were all females and the wings were not in a condition to adequately observe wing venation, Marlíer (1964) declined to provide a species name. Later, Flint (1971Flint ( , 1974 described 2 new Antoptila species from the Amazon and southeastern Brazil. Angrisano (1993) described the female, larvae and pupae of A. brasiliana Mosely, 1939 and males and females of A. plaumanni Flint, 1974 and another new species A. guarani Angrisano, 1993. Subsequently, based on similarities in wing venation and of cases and larval morphology, Flint et al. (1999) synonymized Itauara with Antoptila Mosely 1939, designated A. brasiliana as the type species, and transferred the 3 other known species of Antoptila to Itauara.
In a recent phylogenetic analysis of the entire protoptiline subfamily, Itauara was recovered as a monophyletic group with strong support (Robertson 2010). The presence of a dorsal sheath-like phallicata was identified as a unique synapomorphy of Itauara (Robertson 2010). Although members of this genus have superficially similar male genitalia, certain structures are not homologous. For example, the type species I. brasiliana (Mosely, 1939) has 2 pairs of curious elongate, seta-like processes on sternum IX. The 3 additional species placed in the genus have similarly looking elongate ventral processes, yet they are not the same as in the type species; these processes are parameres, and arise directly from the phallobase or endotheca, rather than sternum IX. The genus Itauara can be identified by features of the male genitalia. The phallic apparatus consists of a sclerotized dorsal sheath covering a very membranous ventral portion, an apparent posterior extension of the phallobase or phallicata. Rarely, the phallicata is tubular or separated from the phallobase by a membranous portion. In some species, this sclerotized dorsal sheath seems to detach from the ventral membrane apically to reveal a single dorsomesal process or spine (e.g., I. amazonica). Mortoniella has a similar dorsomesal process or spine, but in Mortoniella it arises internally from the phallobase, whereas in Itauara it arises dorsobasally, as an extension of the phallicata. In several species the sheath produces a dorsolateral flange-like process, although this character is not diagnostic for the genus. Another genitalic feature characteristic of Itauara is an extremely reduced phallobase. In most species, the phallobase is barely visible, consisting of a small, very lightly sclerotized or an entirely membranous structure. The genera Mastigoptila and Canoptila display similar reductions or absences of the phallobase, but can easily be separated from Itauara by other genitalic characters: Mastigoptila has an elongate, whip-like process arising from the membranes of the phallocrypt; Canoptila has highly membranous digitate parameres. When present (they have been lost in many species), the inferior appendages are rather distinct for Itauara, consisting of a single or apically bifid process produced mesally and fused to the phallobase ventrobasally. This inferior appendage process articulates with the base of the phallobase and in doing so, is capable of pivoting downward ( Fig. 13A and inset). All species, except in I. brasiliana, have rather elongate, sclerotized, rod-like parameres, whose shape varies greatly among species. In many species these parameres arise ventrobasally from the phallobase, with which they appear to articulate. As the inferior appendage process is absent in those species, it is possible that the parameres have taken on a clasper-like function. The forewing venation of Itauara is most similar to that of Cariboptila and Canoptila, with apical forks I-III and a lack of 3A ( Fig. 2A, B). A single species also possesses apical fork IV (Fig. 2C). Canoptila can be differentiated from Itauara by having stout setae occurring below Cu2 whereas in Itauara the setae occur along the vein. Cariboptila can be differentiated from Itauara by the presence of a short discoidal cell, that of Itauara being long. The lengths of the apical forks vary among species. The hind wing venation of Itauara is variable, with either apical forks II, III, and V (Fig. 3C); II and V (Fig. 3A); III only, or II only (Fig. 3B).

Genus
Adult. Body, wings, and appendages pale or tawny brown, often intermingled with rufous or golden hairs, tibia and tarsi yellowish brown (Fig. 1). Wings often with partial white transverse line along anastomosis not reaching costal margin, or often with conspicuous white spot at the arculus (Fig. 1). Head broader than long, vertex rounded, with pair of small anteromesal setal warts or with large anteromesal setal wart, either 1 distinct pair or 1 divided pair of suboval anterior setal warts, small or large suboval posterior warts, suboval or triangular and bulging posterolateral setal warts. Ocelli present. Antennal scape less than or equal to 2 times the length of pedicel. Maxillary palps 5 segmented, 1st and 2nd segments short; 2nd segment bulbous; last 3 segments each nearly same length as 1st and 2nd segments combined. Prothorax with 2 large subtriangular or suboval pronotal setal warts. Mesothorax wider than long, without apparent tegular glands; mesoscutum with pair of suboval anteromesal setal warts, suboval posterolateral warts; mesoscutellum sparsely setose, without distinct setal warts. Forewing (Fig. 2) usually relatively narrow, with margins nearly parallel, occasionally narrowed past anastomosis or much reduced, apex acute, subacute, or rounded. Male occasionally with callosity present in apical costal region of forewing. Forewing venation incomplete, with apical forks I, II, and III present, or rarely I-IV present; Sc and R1 distinct along their entire lengths; fork I sessile or only slightly petiolate with extremely short stem; fork II petiolate or sessile, when petiolate, stem length variable; fork III petiolate, stem variable in length; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing (Fig. 3) margins nearly parallel, tapering only slightly past anastomosis, or narrowed, scalloped past anastomosis, or much reduced; venation variable, either with apical forks II, III, and V present, II and V present, III present, or II present; Sc and R1 fused basally or converging near wing margin; A2 absent. Tibial spurs 1,4,4, rarely 1,3,4, foretibial spur extremely reduced and hairlike. Sixth sternal process present, short and digitate or thumb-like and prominent, apex rounded or attenuate and pointed, usually associated with oblique apodeme posteriorly.
Male genitalia. Segment IX usually rather broad, anterior margin rounded, posterolateral margin without lateral process or lobes in lateral view; tergum IX usually not well developed, simple, and without processes; sternum IX without modification, except in I. brasiliana, which bears 2 pairs of elongate, seta-like processes. Tergum X incompletely fused to tergum IX ventrolaterally or rarely (I. amazonica) completely fused and indistinct from tergum IX, shape extremely variable; dorsomesal margin may be simple without processes, bifid apicomesally, with a single broad, plate-like process, or irregular with several small processes; dorsolateral margin either a simple structure without processes, or more commonly with small paired lobes, elongate, down-turned, finger-like process, or irregular setose processes; ventrolateral margin with paired elongate or broad flange-like processes directed ventrally and sometimes anteriorly, or with one or more irregular, paired, setose, digitate lobes directed posteriorly. Inferior appendages either present or absent; when present, consisting of single or apically bifid process produced mesally, broadest at base and fused to phallobase ventrobasally. Parameres present except in I. brasiliana, arising either ventrobasally from phallobase or laterally from endotheca, sclerotized, shape variable. Phallobase extremely reduced and difficult to discern. Phallicata a sclerotized dorsal sheath covering membranous ventral portion, sometimes receding to a single dorsomesal process arising dorsobasally from phallobase, phallicata occasionally with dorsolateral flange, or occasionally with dorsomesal spine arising posteriorly to phallobase. Endophallus highly membranous, enlarged and convoluted when evaginated, occasionally bearing apical spine-like sclerites and processes.
Female genitalia. (Females unknown for many species.) Truncate posteriorly, not extensible. Abdominal segment VIII short, synscleritous, posterolateral margin slightly incised. Segments IX and X closely associated, with pair of small digitate cerci dorsolaterally.

species relationships
The 22 species of Itauara fall into 2 broad species groups (Table 1). Members of the amazonica species group are recognized by the presence of a fused inferior appendage process. The group is also characterized by the position of the parameres, arising laterally from the endotheca in this group. Species included in the brasiliana species group have completely lost the inferior appendages. When parameres are present (they are vestigial in I. brasiliana) they arise ventrobasally from the phallobase, to which they are often fused. Several members of the brasiliana group also have lateral flange-like processes on the phallicata.
the inferior appendage process of I. alexanderi is broader than that of I. stella. Itauara lucinda differs from I. alexanderi in having a forked paramere and an inferior appendage process that is not bifid. Itauara alexanderi can be distinguished from I. emilia based on differences in the shape of the dorsomesal process of tergum X.
Adult. Body, wings, and appendages fuscous, intermingled with rufous or golden hairs, tibia and tarsi tawny brown. Wings with white transverse line along anastomosis. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem about the same length as fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, associated with oblique apodeme posteriorly.

Itauara amazonica
Description. This species is distinct in having a very elongate inferior appendage process, a rather simple tergum X, and sharply bent apical spines in the endophallus. Itauara amazonica is most similar to 3 species from Guyana, I. bidentata sp. n., I. spiralis sp. n., and I. unidentata sp. n. These species, including I. amazonica, all have a dorsomesal spine arising from the phallicata. However, in I. amazonica, this spine appears as a short, posterior extension of the phallicata, whereas in the other species, the spine arises basally, as a separate spine. I. amazonica can further be distinguished from these species based on differences in the shape of tergum X and parameres. Adult. Body, wings, and appendages pale or tawny brown in alcohol. Forewing relatively narrow, with margins nearly parallel, apex subacute. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem shorter than fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing narrow and slightly scalloped past anastomosis; apical fork III present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process short and digitate, apex attenuate and pointed, associated with weak oblique apodeme posteriorly.
Male genitalia. Preanal appendages absent. Segment IX relatively broad; anterior margin rounded; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X completely fused to tergum IX, divided or bifid apicomesally, each half terminating in pointed process directed posteriorly; dorsolateral margin without processes; ventrolateral margin without processes. Inferior appendages present as single, elongate setose process produced mesally, broadest at base and fused to phallobase ventrobasally. Parameres present, paired, inserted in membranous lobe, arising laterally from endotheca, sclerotized and rod-like, slender and elongate, upturned, directed dorsally, apex pointed. Phallobase reduced, lightly sclerotized. Phallicata forming a long sclerotized dorsal sheath extending from phallobase, bent sharply upward at middle, divided apicomesally, terminating in 2 pointed processes. Endophallus membranous, enlarged and convoluted when invaginated, receding anterior to apex of phallicata, ventrally bearing a pair of broad, tooth-like downturned processes, apically with pair of sharply bent sclerotized spines, pointing anteriorly.
Paratypes: BRAZIL: Amazonas: same data as holotype -3 males, 2 females (NMNH). Description. Itauara bidentata can be diagnosed by its large, bifid paramere process, and spade-like shaped inferior appendage. It is most similar to I. unidentata sp. n., which has a similarly shaped tergum X, dorsomesal spine, and apical sclerites. The 2 species can be separated by their paramere processes; in I. unidentata the paramere consists of a single large tooth-like spine, whereas in I. bidentata, the paramere process is bifid. Itauara spiralis, sp. n., has a similarly shaped tergum X, but is easily distinguished from I. bidentata by differences in the shape of the inferior appendage process, parameres, and phallicata. Adult. Body, wings, and appendages pale or tawny brown in alcohol. Forewing relatively narrow, with margins nearly parallel, apex subacute. Forewing venation incomplete, with apical forks I, II, and III present; fork I sessile; fork II petiolate, stem about the same length as fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing narrow and slightly scalloped past anastomosis; apical fork II present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process short and digitate, apex attenuate and pointed, associated with strong oblique apodeme posteriorly.

Itauara blahniki
Adult. Body, wings, and appendages pale or tawny brown, often intermingled with rufous or golden hairs, tibia and tarsi tawny brown. Wings with conspicuous white spot at the arculus. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II sessile; fork III petiolate, stem about the same length as fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II, III, and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, associated with oblique apodeme posteriorly.
Etymology. We are delighted to name this species for Dr. Roger Blahnik, in honor of his many contributions to the systematics of Protoptilinae.

Itauara brasiliana (Mosely 1939)
http://species-id.net/wiki/Itauara_brasiliana Fig. 2A Description. This species is easily diagnosed by the presence of 2 pairs of extremely elongate, seta-like processes on sternum IX. Itauara brasiliana is also distinct in having vestigial parameres, consisting only of very small, setose lobes. Another distinguishing characteristic is the shape of segment IX, which is rather narrow, and receded ventrally. The rather elongate profile of tergum X, as well as the shape of the dorsomesal Figure 8. Itauara brasiliana (Mosely, 1939). Male genitalia A lateral B dorsal C ventral. Abbreviations: enph. = endophallus; phc. = phallicata; pmr . = paramere; t. X = tergum X; vl. pr. = ventrolateral process. and lateral margins of tergum X, slightly resembles that of I. plaumanni (Flint 1974). However, I. plaumanni has much more pronounced, elongate parameres, and lacks the seta-like processes on sternum IX. Adult. Body, wings, and appendages pale or tawny brown in alcohol. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem shorter than fork; fork III petiolate, stem about the same length as fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II and V present; Sc and R1 converging near wing margin; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, associated with oblique apodeme posteriorly.
Male genitalia. Preanal and inferior appendages absent. Segment IX ventrally narrow, broad medially; anterior margin relatively straight from dorsum to medial area, ventral portion rounded; posterolateral margin highly membranous, receding ventrally; sternum IX bearing 2 pairs of extremely elongate, seta-like processes. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin subtriangular, slightly upturned; dorsolateral margin slightly irregular, without processes; ventrolateral margin with 2 pairs of processes, the upper an elongate finger-like process slightly downturned, the lower a smaller lobe-like setose process. Parameres vestigial, consisting of a pair of small, digitate setose lobes arising ventrolaterally from endotheca. Phallobase apparently absent or entirely membranous. Phallicata forming a long, lightly sclerotized dorsal sheath, sinuous, broadest medially, narrowed distally. Endophallus membranous, enlarged and convoluted when invaginated. Description. This species is only known from the male holotype. Itauara charlotta is diagnosed by the unique combination of several male genitalic characters. The dorsomesal margin of tergum X has several small, irregular, setose processes. Itauara tusci sp. n., has a similarly shaped dorsomesal margin, but differs in the shape of the parameres and phallica- ta. The parameres of I. charlotta are similar to those of I. flinti sp. n.; both arise ventrobasally from the phallobase, and are curved and downturned. Itauara flinti is easily separated from I. charlotta based on differences in the shape of tergum X and phallicata processes.
Adult. Body, wings, and appendages pale or tawny brown in alcohol. Wings with conspicuous white spot at the arculus and faint transverse line along anastomosis. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem shorter than fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II, III, and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, often associated with weak oblique apodeme posteriorly.
Material examined. Holotype male: BRAZIL: Minas Gerais: Serra do Cipó, Cardeal Mota, Cachoeira Veu da Noiva, 19°18'55"S, 043°36'16"W, 800 m, 12.11.2001 (Holzenthal, Amar., Blahnik, Paprocki) Etymology. We are delighted to name this species for the senior author's mother, Charlotte Ruth Robertson. Description. This species is known only from the male holotype. Itauara emilia can be recognized by the distinct, rather blunt shape of the dorsomesal process of tergum X. The species is similar to I. alexanderi sp. n., I. lucinda sp. n., and I. stella sp. n., as discussed under each of those species. Each of these species possess an inferior appendage process, a dorsomesal process on tergum X, and rather sinuous parameres. Itauara emilia is most similar to I. alexanderi and I. stella in having similarly shaped parameres and an apically bifid inferior appendage process. Itauara emilia differs from these 2 species in having a much more elongate inferior appendage process and a blunt dorsomesal process on tergum X.
Adult. Body, wings, and appendages tawny brown (specimen missing hairs). Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem about the same length as fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, associated with oblique apodeme posteriorly.

Itauara flinti Robertson & Holzenthal, sp. n.
urn:lsid:zoobank.org:act:08207511-83D1-46A4-8168-BAE2935AAC0A http://species-id.net/wiki/Itauara_flinti Fig. 11A-C Description. This species is known only from the male holotype. It is diagnosed by a unique combination of male genitalic characters and can be recognized by the shape of the phallicata process. Itauara flinti is most similar to I. charlotta sp. n., by having similarly shaped parameres, but these species are easily separated based on differences in the shape of tergum X and the phallicata process. The elongate and downturned dorsomesal process of tergum X is similar to that of I. guarani (Angrisano 1993), but the 2 species differ in the shape of the parameres and phallicata processes. The ventrolateral process of tergum X is similar to that of I. tusci sp. n. These species differ in the shape of the parameres, dorsomesal margins of tergum X, and phallicata processes.
Adult. Body, wings, and appendages pale or tawny brown in alcohol. Forewing relatively narrow, with margins nearly parallel, apex subacute. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I petiolate, but with extremely short stem; fork II petiolate, stem about the same length as fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing narrow and slightly scalloped past anastomosis; apical forks II and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, associated with weak oblique apodeme posteriorly.
Male genitalia. Preanal and inferior appendages absent. Segment IX dorsally narrow, broad ventrally; anterior margin rounded; posterolateral margin lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin with single, downturned, elongate process; dorsolateral margin irregular and setose; ventrolateral margin with 2 pairs of processes, the upper a small lobe-like setose process, the lower an elongate finger-like process bearing a few elongate setae. Parameres present, paired, arising ventrobasally from fused endotheca and phallobase, sclerotized and rod-like, slender and elongate, downturned, curved basally, straight medially and distally, directed ventrally and inward, apex pointed. Phallobase reduced, lightly sclerotized. Phallicata forming a long, lightly sclerotized dorsal sheath, slightly sinuous, medially with pair of lightly sclerotized rounded lateral flanges projecting posteroventrally, ventrally with several sclerotized points, thorn-like apices directed inward. Endophallus membranous, enlarged and convoluted when invaginated, with 1 large upper lobe and 1 smaller lower lobe.

Etymology.
We are delighted to name this species for Dr. Oliver Flint, Jr. in honor of his numerous important contributions to Neotropical caddisfly taxonomy. (Angrisano, 1993) http://species-id.net/wiki/Itauara_guarani Fig. 12A Description. This species can be recognized by the very broad, lateral flanges on the phallicata. The phallicata of I. guarani is slightly sinuous, with a lightly sclerotized basal portion and a rugous or almost membranous distal portion. Itauara simplex sp. n., also has a very lightly sclerotized phallicata, but the 2 species differ in the shape of the parameres and I. simplex lacks the processes of phallicata. The parameres of I. guarani arise ventrobasally from the phallobase and are sinuous, like those in I. plaumanni. However, the phallicata in I. plaumanni is much more sclerotized and the 2 species also differ in the shape of tergum X and the phallicata processes.

Itauara guarani
Adult. Body, wings, and appendages pale or tawny brown in alcohol. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I petiolate, but with extremely short stem; fork II petiolate, stem shorter than fork; fork III petiolate, stem longer than fork; Cu1 incomplete, not reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II and III present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process short and digitate, apex rounded, associated with weak oblique apodeme posteriorly.
Male genitalia. Preanal and inferior appendages absent. Segment IX dorsally narrow, broad ventrally; anterior margin rounded; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin with single, downturned, elongate process; dorsolateral margin irregular and setose; ventrolateral margin with paired subtriangular setose process directed posteriorly. Parameres present, paired, arising ventrobasally and fused to phallobase, sclerotized and rod-like, slender and elongate, sinuous, strongly downturned basally, distal portion slightly upturned and broader, directed posteriorly, apex pointed. Phallobase reduced, lightly sclerotized. Phallicata sinuous, with lightly sclerotized base, distal portion membranous, with pair of broad, sclerotized wing-like lateral flanges.   Fig. 2B, 3B, 13A-C Description. Itauara guyanensis has distinct extremely sinuous, almost corkscrewshaped, parameres. The phallicata is short, sclerotized, and upturned apically and with 2 pointed lateral processes. The species can also be recognized by the thumb-like shape of the inferior appendage process and the presence of a bifid apical process in the endophallus. I. guyanensis is somewhat similar to I. jamesii sp. n., and resembles that species in the shape of the inferior appendage process and sinuous parameres. The 2 species can be easily separated by differences in the shape of tergum X and the phallicata. The species I. alexanderi sp. n., I. emilia sp. n., and I. stella sp. n., also have rather sinuous parameres, but differ in the shape of the phallicata, tergum X, and several other characters.
Adult. Body, wings, and appendages pale or tawny brown in alcohol. Forewing narrow past anastomosis, apex acute. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem about the same length as fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing narrow and slightly scalloped past anastomosis; apical fork II present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,3,4, foretibial spur extremely reduced and hairlike. Sixth sternal process short and digitate, apex attenuate and pointed, associated with strong oblique apodeme posteriorly.
Etymology. This species is named for the country of Guyana, where the specimens were collected.

Itauara jamesii Robertson & Holzenthal, sp. n.
urn:lsid:zoobank.org:act:9EAE24A5-16C2-4B9F-8BDD-42976CE0E569 http://species-id.net/wiki/Itauara_jamesii Fig. 14A-C Description. Itauara jamesii is known only from the male holotype, and its relationship to other species in not immediately evident. The parameres have a rather asymmetrical aspect, but it is possible that this particular specimen is distorted. The species has an inferior appendage process like several other species, but has a distinct subtriangular shape. The dorsomesal margin of tergum X is bifid, each half a small setose protuberance. Itauara peruensis sp. n., also has a bifid dorsomesal margin, but in that species, it appears as a prominent process with pointed apices.
Adult. Body, wings, and appendages pale or tawny brown, often intermingled with rufous or golden hairs, tibia and tarsi tawny brown. Wings with conspicuous white spot at the arculus and faint transverse line along anastomosis. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex subacute. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem shorter than fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, associated with weak oblique apodeme posteriorly. Male genitalia. Preanal and inferior appendages absent. Segment IX dorsally and ventrally narrow, broad medially; anterior margin rounded; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin bifid and slightly produced, each half small, setose, with a rounded apex; dorsolateral margin without processes; ventrolateral margin with paired, broad, semi-circular setose flange-like process. Inferior appendages present as single, broad, subtriangular setose process, fused to phallobase ventrobasally, bearing small digitate lobes ventrolaterally, each bearing a seta. Parameres present, paired, inserted in membranous lobe, arising laterally from endotheca, sclerotized and rod-like, extremely sinuous, seemingly asymmetrical, apex pointed. Phallobase reduced, lightly sclerotized with phallic shield. Phallicata forming a long sclerotized dorsal sheath extending from phallobase, broadest basally, bent upward at middle, with paired sclerotized concave discs arising basodorsally and forming a connection with posterior margin of segment IX. Endophallus membranous, enlarged and convoluted when invaginated.

Itauara julia Robertson & Holzenthal, sp. n.
urn:lsid:zoobank.org:act:5DE975A9-37FA-4BB0-9A52-662621648658 http://species-id.net/wiki/Itauara_julia Fig. 1, 3C,15A-C Description. This species is distinct in having a curved, spatulate ventral process in the endophallus and having a phallicata that is not continuous with the phallobase. The parameres of I. julia are strongly bent dorsally at the base, and terminate in a sharp, downturned point. The parameres curve upward, are fused to and continuous with the phallobase, and arise ventrobasally. The parameres of I. blahniki sp. n., I. rodmani sp. n., and I. tusci sp. n., are similarly structured. However, in these species, the parameres are not as abruptly bent.
Adult. Body, wings, and appendages pale or tawny brown, often intermingled with rufous or golden hairs, tibia and tarsi tawny brown. Wings with conspicuous white spot at the arculus. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. With apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II sessile; fork III petiolate, stem shorter than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II, III, and V pre- sent; Sc and R1 converging near wing margin; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex attenuate and pointed, associated with weak oblique apodeme posteriorly.
Male genitalia. Preanal and inferior appendages absent. Segment IX dorsally narrow, broad ventrally; anterior margin relatively straight from dorsum to medial area, ventral portion blunt; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin straight, without processes; dorsolateral margin with several small irregular setose processes; ventrolateral margin with paired subquadrate setose process projecting ventrally. Parameres present, paired, arising ventrobasally and fused to phallobase, sclerotized and rod-like, slender and elongate, strongly bent upward basally, apex pointed and downturned, ventrobasally with a small patch of setae. Phallobase reduced, lightly sclerotized. Phallicata forming a long sclerotized dorsal sheath, mostly straight, broadest basally, apex with apicomesal point and 2 lateral downturned points. Endophallus membranous, enlarged and convoluted when invaginated, ventrally with a curved spatulate process.
Adult. Body, wings, and appendages fuscous, intermingled with rufous or golden hairs, tibia and tarsi tawny brown. Wings with white transverse line along anastomo- sis. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex subacute. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem about the same length as fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex attenuate and pointed, associated with oblique apodeme posteriorly.
Male genitalia. Preanal appendages absent. Segment IX dorsally narrow, broad medially and ventrally; anterior margin rounded; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin with single, downturned, elongate process; dorsolateral margin without processes; ventrolateral margin with paired, broad flange-like setose process with small upper lobe and larger subquadrate lower lobe. Inferior appendages present as single, broad, subquadrate setose process, broadest basally, fused to phallobase ventrobasally, with 2 pairs of small digitate lobes ventrolaterally, each bearing a seta. Parameres present, paired, inserted in membranous lobe, arising laterally from endotheca, sclerotized and rod-like, bifid, with short lower process and longer, slightly medially bent upper process, directed posteriorly, apices pointed. Phallobase reduced, lightly sclerotized. Phallicata forming a long, straight sclerotized dorsal sheath extending from phallobase. Endophallus membranous, enlarged and convoluted when invaginated, with 1 upper and 1 lower lobe. Perhaps the most notable feature of this species is the extremely curved, ram-like shaped parameres. Itauara spiralis sp. n., also has highly curved, spiral shaped parameres, but in I. spiralis, the paramere is curved along the entire length of the paramere, whereas in I. ovis, the paramere is curved basally, but straight distally. The 2 species differ in other respects, including the shape of the inferior appendage process. In I. ovis, the inferior appendage process is elongate, and rather inflated apically; in I. spiralis, the inferior appendage process is bifid. The 2 species also differ in the shape of tergum X. In I. ovis, tergum X is slightly notched apicomesally and has 2 pairs of rather small, subtriangular ventrolateral processes. In I. spiralis, tergum X is not notched, but has a pair of very long, finger-like dorsomesal processes and a pair of very broad, irregular ventrolateral processes. The phallicata of I. ovis is also quite distinct, being rather broad, and saddle-shaped, with a dorsobasal hump and upturned apex.
Adult. Body, wings, and appendages fuscous, intermingled with rufous or golden hairs, tibia and tarsi tawny brown. Wings often with a few pale cream-colored or white hairs at arculus. Forewing relatively narrow, with margins nearly parallel, apex subacute. Forewing venation incomplete, with apical forks I, II, and III present; fork I sessile; fork II petiolate, stem about the same length as fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing narrow and slightly scalloped past anastomosis; apical forks II and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process short and digitate, apex attenuate and pointed, associated with strong oblique apodeme posteriorly.
Male genitalia. Preanal appendages absent. Segment IX ventrally narrow, broad medially; anterior margin rounded; posterolateral margin lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin subtriangular, slightly produced with small cleft; dorsolateral margin without processes; ventrolateral margin with an outer pair of subtriangular setose processes directed ventrally, and an inner pair of subtriangular processes directed posteroventrally. Inferior appendages present as single, rather elongate setose process produced mesally, apex broad and slightly irregular, fused to phallobase ventrobasally. Parameres present, paired, inserted in membranous lobe, arising laterally from endotheca, sclerotized and rod-like, ram-like, curving 360 degrees at base, distal portion straight, directed posteriorly, apex pointed. Phallobase extremely reduced and difficult to discern. Phallicata forming a saddle-shaped sclerotized dorsal sheath, with dorsal hump basally, distal portion curving upward. Endophallus membranous, enlarged and convoluted when evaginated, with pair of elongate lateral sclerites ventrally. Etymology. The name ovis, comes from the Latin for sheep, and is suggested by the shape of the parameres, which are reminiscent of a ram's horn.

Itauara peruensis Robertson & Holzenthal, sp. n.
urn:lsid:zoobank.org:act:0F3E4131-B78D-470A-9DA5-83228D16BEC6 http://species-id.net/wiki/Itauara_peruensis Fig. 18A-C Description. This species is distinct in having a lightly sclerotized endophallus. Ventrally, the endophallus is membranous, but in lateral view, it has the appearance of being entirely sclerotized. Another unique feature is the prominent bifid dorsomesal process of tergum X. Itauara peruensis has a rather elongate inferior appendage process like I. ovis, but it is not inflated apically like that species. The species also differ in the shape of the parameres; those of I. peruensis are nearly straight, while those of I. ovis are spiral-shaped.
Adult. Body, wings, and appendages pale or tawny brown in alcohol. Forewing narrow past anastomosis, apex acute. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem about the same length as fork; fork III petiolate, stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing narrow and slightly scalloped past anastomosis; apical fork II present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process short and digitate, apex attenuate and pointed, associated with strong oblique apodeme posteriorly.

Itauara plaumanni
Description. Itauara plaumanni can be recognized by the irregular lobe-like shape of the lateral flanges on the phallicata. Additionally, this species has a rather elongate segment IX and tergum X. The profile of tergum X resembles that of I. brasiliana, but I. plaumanni has fewer lateral processes than that species. Itauara plaumanni has parameres similar in shape to those of I. guarani, yet these species are easily separated by differences in the shape of the lateral flanges of the phallicata, and tergum X. Adult. Body, wings, and appendages pale or tawny brown in alcohol. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem shorter than fork; fork III petiolate, stem about the same length as fork; Cu1 complete, reaching Figure 19. Itauara plaumanni (Flint, 1974) (UMSP000086359). Male genitalia A Sternum VI process; ttt B lateral C dorsal D ventral. Abbreviations: dl. pr. = dorsolateral process; enph. = endophallus; phb = phallobase; phc. = phallicata; phc. pr. = phallicata process; pmr . = paramere; t. X = tergum X. wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II, III, and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process short and digitate, apex attenuate and pointed.
Male genitalia. Preanal and inferior appendages absent. Segment IX relatively broad; anterior margin rounded; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin slightly produced as small irregular point; dorsolateral margin with 2 pairs of processes, the upper an elongate subtriangular process slightly downturned, the lower a small lobe-like setose process; ventrolateral margin without processes. Parameres present, paired, arising ventrobasally and fused to phallobase, sclerotized and rod-like, slender and elongate, sinuous, downturned basally, distal portion slightly upturned, directed posteriorly and inward, apex pointed, ventrobasally with small patch of setae. Phallobase reduced, lightly sclerotized. Phallicata forming a long sclerotized dorsal sheath, slightly bent upward medially, with pair of irregular lobe-like lateral flanges projecting posteroventrally. Endophallus membranous, enlarged and convoluted when invaginated, with lightly sclerotized lobe ventrally and laterally, containing 2 small sclerites. Description. This species is very similar to I. tusci sp. n., which also has very long, upturned, tusk-like parameres and a strongly upturned phallicata. The 2 species can be inset, variant C dorsal D ventral; inset, tergum X. Abbreviations: dl. pr. = dorsolateral process; enph. = endophallus; phb = phallobase; phc. = phallicata; phc. pr. = phallicata process; pmr . = paramere; t. X = tergum X; vl. pr. = ventrolateral process. separated based on the shape of the phallicata process, which is pointed and blade-like in I. tusci and rounded or blunt in I. rodmani. The phallicata is also more sclerotized in I. rodmani. Additionally, the dorsomesal margin of tergum X is irregular, with several small setose processes in I. tusci, whereas in I. rodmani, the dorsomesal margin is rather smooth and triangular. The ventrolateral processes of the 2 species also differ: I. tusci has a small upper and more elongate lower process; I. rodmani has a single, short, digitate process. I. blahniki also has upturned, tusk-like parameres, but is easily distinguished from I. rodmani based on differences in tergum X and the phallicata. Itauara blahniki and I. blahniki, sp. n., also has tusk-like parameres, but they are not as long and curved as I. rodmani.
Adult. Body, wings, and appendages pale or tawny brown in alcohol. Wings with conspicuous white spot at the arculus. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex subacute. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem shorter than fork; fork III petiolate, stem about the same length as fork; Cu1 incomplete, not reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II, III, and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, associated with weak oblique apodeme posteriorly.
Male genitalia. Preanal and inferior appendages absent. Segment IX relatively broad; anterior margin rounded; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin subtriangular, very slightly downturned; dorsolateral margin with paired small, slightly down-turned, setose process; ventrolateral margin with an outer and inner pair of small setose processes directed posteriorly. Parameres present, paired, arising ventrobasally from fused endotheca and phallobase, sclerotized and rod-like, tusklike, strongly curving upward, apex pointed. Phallobase reduced, lightly sclerotized with phallic shield. Phallicata forming a long sclerotized dorsal sheath extending from phallobase, strongly curving upward with apex directed dorsally, with pair of broad, sclerotized wing-like lateral flanges with rounded or subquadrate ventral margins. Endophallus membranous, enlarged and convoluted when invaginated, with 1 tubular upper lobe and 3 smaller lower lobes.
Etymology. We are delighted to name this species for Dr. James Rodman, the NSF program director who initiated the Partnership for Enhancing Expertise in Taxonomy (PEET) program. The PEET program provides funding for the training of taxonomists of little known organisms. The senior author is grateful for the wonderful experience she had while participating in the PEET program as a doctoral student and the opportunity to study Trichoptera taxonomy. Description. This species can be recognized by its rather simple genitalic capsule. Tergum X is produced dorsomesally into a broad, elongate plate and has just one small ventrolateral process. The parameres are relatively short, straight basally, but slightly bent downward distally. The phallicata is short and very lightly sclerotized, and the endophallus is large and membranous, with 2 lateral patches or elongate setae apically. Itauara guarani also has a very lightly sclerotized phallicata, but the 2 species differ in the shape of the parameres and I. guarani has lateral flange-like processes on the phallicata.

Itauara simplex
Adult. Body, wings, and appendages fuscous, intermingled with rufous or golden hairs. Wings with conspicuous white spot at the arculus. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex subacute. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I petiolate, but with extremely short stem; fork II petiolate, stem about the same length as fork; fork III petiolate; stem longer than fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical fork II present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex rounded, associated with weak oblique apodeme posteriorly.
Paratype: BRAZIL: São Paulo: same data as holotype -1 female (UMSP). Etymology. This species is so named for the rather simple structure of the phallic apparatus and genital capsule.
Itauara spiralis Robertson & Holzenthal, sp. n. urn:lsid:zoobank.org:act:0568AD89-DD46-4FCB-87F4-0F9D884AEC04 http://species-id.net/wiki/Itauara_spiralis Fig. 22A-C Description. This species is distinct in having a sclerotized, tubular phallicata, and an elongate, laterally compressed, dorsomesal spine. The phallicata in other species are less tubular, appearing as a dorsal sheath. This dorsal sheath was identified as a synapomorphy for the genus in a previous phylogenetic study of Protoptilinae (see Chapter 1, this work). Itauara spiralis was not included in that study, however, I. spiralis is placed in Itauara since it shares many other characteristics common to the genus such as an inferior appendage process, and a tergum X that is nearly identical to I. bidentata sp. n. and I. unidentata sp. n.
Itauara spiralis can be recognized by the extremely curved, spiral-shaped parameres. I. ovis, sp. n., also has highly curved, spiral shaped parameres, but in I. spiralis, the paramere is curved along the entire length of the paramere, whereas in I. ovis, the paramere is curved basally, but straight distally. The 2 species differ in other respects, including the shape of the inferior appendage process, which is bifid in I. spiralis and inflated apically in I. ovis. The 2 species also differ in the shape of tergum X and the phallicata. Tergum X is very similar to those of I. bidentata and I. unidentata; all have elongate, finger-like dorsolateral processes and broad, irregular, setose ventrolateral processes. Itauara spiralis is distinguished from these other 2 species by having a bifid inferior appendage process, spiral-shaped parameres, and laterally compressed dorsomesal spine.
Adult. The only specimen of this species is in very poor condition. Therefore, head, thoracic, and wing characters could not be observed. However, the genitalia are intact.
Male genitalia. Preanal appendages absent. Segment IX ventrally narrow, broad medially; anterior margin rounded; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin straight, shallowly excavate; dorsolateral margin with paired elongate, down-turned, finger-like process; ventrolateral margin with paired, very broad flange-like setose process consisting of several small irregular lobes. Inferior appendages present as apically bifid, setose process produced mesally, broadest at base and fused to phallobase ventrobasally. Parameres present, paired, arising laterally from endotheca, spiral-shaped, curving 360 degrees at base with curve continuing to apex, directed posteroventrally, apex pointed. Phallobase reduced, lightly sclerotized. Phallicata forming a short slerotized dorsal tube extending from phallobase, with a long, broad dorsomesal spine arising posteriorly to phallobase. Endophallus membranous, rather small, apically sharply bent downward, pointing anteroventrally. Description. This species is associated with Itauara alexanderi sp. n., I. emilia sp. n., and I. lucinda sp. n., as discussed under each of those species. Each of these species possess an inferior appendage process, a dorsomesal process on tergum X, and rather sinuous parameres. Of these species, I. stella is most similar to I. alexanderi. Both of these species have similarly shaped elongate dorsomesal processes and broad ventrolateral processes of tergum X. Both also have apically bifid inferior appendage processes. Itauara stella can be distinguished from by the length of the parameres; those of I. stella are longer and more strongly directed laterally than those of I. alexanderi. Additionally, the inferior appendage process of I. alexanderi is broader than that of I. stella. Itauara stella can be differentiated from I. emilia by the shape of the dorsomesal process and from I. lucinda by the shape of the parameres and inferior appendage process. Adult. Body, wings, and appendages pale or tawny brown in alcohol. Wings with white transverse line along anastomosis. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; stem about the same length as fork; fork III petiolate, stem about the same length as fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumb-like, apex attenuate and pointed, associated with weak oblique apodeme posteriorly.
Etymology. We are delighted to name this species for the senior author's daughter, Stella Claire Thompson. Description. This species is very similar to I. rodmani sp. n., which also has very long, upturned, tusk-like parameres and a strongly upturned phallicata. The 2 species are separated based on the shape of the phallicata process, which is pointed and blade-like in I. tusci and rounded or blunt in I. rodmani. The phallicata of I. tusci is more lightly sclerotized than I. rodmani, especially at the distal portion. Additionally, in I. tusci, the dorsomesal margin of tergum X is irregular, with several small setose processes, whereas in I. rodmani, the dorsomesal margin is rather smooth and triangular. The ventrolateral processes of the 2 species also differ: I. tusci has a small upper and more elongate lower process; I. rodmani has a single, short, digitate process. Itauara blahniki also has upturned, tusk-like parameres, but is easily distinguished from I. tusci based on differences in tergum X and the phallicata.

Itauara tusci
Adult. Body, wings, and appendages pale or tawny brown, often intermingled with rufous or golden hairs, tibia and tarsi tawny brown. Wings with conspicuous white spot at the arculus. Forewing slightly broader past anastomosis, but with margins nearly parallel, apex rounded. Forewing venation incomplete, with apical forks I, II, and III present; Sc and R1 distinct along their entire lengths; fork I sessile; fork II petiolate, stem shorter than fork; fork III petiolate, stem about the same length as fork; Cu1 complete, reaching wing margin; Cu1 and Cu2 intersecting near anastomosis; row of erect setae present along Cu2; A3 absent; crossveins forming a relatively linear transverse cord; discoidal cell longer than Rs vein. Hind wing margins nearly parallel, tapering only slightly past anastomosis; apical forks II, III, and V present; Sc and R1 fused basally; A2 absent. Tibial spurs 1,4,4, foretibial spur extremely reduced and hairlike. Sixth sternal process thumblike, apex attenuate and pointed, associated with weak oblique apodeme posteriorly.
Itauara unidentata Robertson & Holzenthal, sp. n. urn:lsid:zoobank.org:act:63403AB7-370C-4DA2-8628-C630D0C9C1E6 http://species-id.net/wiki/Itauara_unidentata Fig. 2C, 25A-C Description. This species can be diagnosed by its large, tooth-like paramere process, and broad inferior appendage process. It is most similar to I. bidentata sp. n., which has a similarly shaped tergum X, dorsomesal spine, and apical sclerites. The 2 species can be separated by their paramere processes; in I. unidentata the paramere consists of a single large tooth-like process, whereas in I. bidentata, the paramere process is bifid. Itauara amazonica also has a dorsomesal spine, but can be distinguished from I. unidentata by the simple shape of tergum X and parameres. Itauara spiralis, sp. n., has a similarly shaped tergum X, but is easily distinguished from I. unidentata by differences in the shape of the inferior appendage process, parameres, and phallicata.
Male genitalia. Preanal appendages absent. Segment IX dorsally and ventrally narrow, broad medially; anterior margin rounded; posterolateral margin membranous or very lightly sclerotized; sternum IX without modification. Tergum X incompletely fused to tergum IX with membrane or lightly sclerotized region ventrolaterally; dorsomesal margin straight, without processes; dorsolateral margin with paired elongate, down-turned, finger-like process; ventrolateral margin with paired, broad flange-like setose process consisting of several small irregular lobes. Inferior appendages present as single, broad, irregular setose process, broadest basally, fused to phallobase ventrobasally, bearing a single pair of small digitate lobes ventrolaterally, each bearing a seta. Parameres present, paired, arising laterally from endotheca, strongly sclerotized, large tooth-like process, curving ventrally and outward, apex pointed. Phallobase reduced, lightly sclerotized dorsally, laterally membranous, with 2 irregular and elongate sclerites arising basolaterally. Phallicata forming a short slerotized dorsal sheath with an elongate dorsomesal spine arising posteriorly to phallobase. Etymology. The name unidentata is suggested by the single tooth-like paramere process.