from Brazil - Biotaxa

Zootaxa 4139 (1): 051–075 http://www.mapress.com/j/zt/ Copyright © 2016 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://doi.org/10.11646/zootaxa.4139.1.3 http://zoobank.org/urn:lsid:zoobank.org:pub:C40885DD-FB4C-461C-92CE-143BF7174E76

Taxonomy of Atopsyche Banks (Trichoptera: Hydrobiosidae) from Brazil: New species, distributional notes and identification key VICTOR GOMES1 & ADOLFO RICARDO CALOR1,2 1

Universidade Federal da Bahia, Instituto de Biologia, Departamento de Zoologia, PPG Diversidade Animal, Laboratório de Entomologia Aquática - LEAq. Rua Barão de Geremoabo, 147, campus Ondina, Ondina, CEP 40170-115, Salvador, Bahia, Brazil. E-mail: [email protected] 2 Corresponding author. E-mail: [email protected]

Abstract Three new species of Atopsyche Banks 1905 are described and illustrated from Brazil: Atopsyche diamantina n. sp., A. kamakan n. sp., and A. muelleri n. sp. New records of A. apurimac Schmid 1989, A. sanctipauli Flint 1974, A. serica Ross 1953, and A. zernyi Flint 1974 are included, as well as the first records to states of Bahia, Goiás, and Rio Grande do Sul. Atopsche rinconi Holzenthal & Cressa 2002 is recorded from Brazil for the first time. An identification key is also provided for species of the genus from Brazil. Moreover additional characters and illustrations of A. sanctipauli and A. zernyi are presented. Key words: aquatic insects, Atlantic forest, Atopsaura, caddisfly, Hydrobiosinae, Neotropical Region

Introduction Atopsyche Banks 1905, with type species Atopsyche tripunctata Banks 1905 (original designation), belongs to Hydrobiosidae Ulmer 1905, initially a subfamily Hydrobiosinae of Rhyacophilidae Stephens 1836. The type genus Hydrobiosis McLachlan 1868 originally belonged to Hydropsychidae Curtis 1835 and was later made type genus of Hydrobiosinae, that includes two other genera, Psilochorema McLachlan 1866 and Philanisus Walker 1852 (Ulmer 1905). Ulmer (1907) further refined the diagnostic characters of the subfamily to include those of Apsilochorema Ulmer 1907 and Atopsyche (Ward et al. 2004). Ross (1951a, 1951b, 1953 as Hydrobiosini; 1956 as Hydrobiosinae) and Ross and King (1951, 1952 as Hydrobiosini) provided a phylogeny of the Hydrobiosini (or Hydrobiosinae) in Rhyacophilidae, showing the monophyletic status of the tribe (or subfamily) based on the following characters: Forewings each with r3-r4 (“s”) crossvein present; forewing anal cells long; forewing vein Cu2 sinuate; male filipods (“filicerci”) present; and larval forelegs chelate. Schmid (1970) revised Rhyacophilidae, elevating Hydrobiosinae to family status, Hydrobiosidae. The family Hydrobiosidae comprises nearly 420 described species in 56 genera (50 extant genera, 6 fossil genera, Morse personal communication) and is recorded from all biogeographical regions, except the Afrotropical region. Hydrobiosidae can be recognized by the presence of ocelli, antennae slightly shorter than forewings, maxillary palps long and slender, spur formula 2,4,4; presence of ventral lobes on male sternites VI and VII and on female sternites V and VI (Schmid 1989). The larvae are free-living predators in running water, and the genus Atopsyche has forelegs modified to form specialized pincer-like structures for securing prey (Reynaga & Rueda Martín 2010). At pupation, they build pupal chambers from small stones and these are anchored to the substrate by silk (Holzenthal et al. 2007). Schmid (1989) revised the family, including a review of genus Atopsyche with more than 45 new species. He criticized the phylogenetic skill of Ross and King (1952) and Ross (1953) (calling Ross a “assez piètre phylogénéticien”), but adopted their classification “without too much criticism,” calling it “approximate, provisional, insufficient, and not without errors” (Schmid 1989). Despite these concerns, we also follow that Accepted by J. Morse: 10 Jun. 2016; published: 19 Jul. 2016

51

classification, in which Atopsyche is divided into three subgenera: Atopsaura Ross 1953, Atopsyche, and Dolochorema Banks 1913. The first two subgenera are divided into diagnostic species groups mainly by the shape of the inferior appendages and the phallic apparatus. The subgenus Atopsaura is composed of three species groups (A. batesi, A. falina and A. longipennis Groups) and Atopsyche of two species groups (A. bolivari and A. tripunctata Groups). Another diagnostic species group (A. bicolorata Group) resembles Dolochorema species, but it is incertae sedis with respect to subgenus; two species are “isolated species,” not assigned to any subgenus or species group; and one species name is a nomen dubium, with unknown or doubtful application. While describing new species from Costa Rica, Blahnik and Gottschalk (1997) commented about the need for an updated phylogenetic framework for the genus that includes the species described since Schmid’s (1989) revision. The Neotropical genus Atopsyche is the most diverse in the family, comprising 132 described species and occurring from southwestern USA to South America, except the Chilean Subregion (Holzenthal & Cressa 2002). The genus can be characterized by long and slender maxillary palps and forewings with vein setae well developed (Schmid 1989). The larva can be recognized by having each foretibia and foretarsus fused and moving against a concave apicoventral extension of the forefemur, producing a chelate foreleg useful for capturing prey (Wiggins 2004). Larvae are free-living, occurring typically in cold and rapid streams. Until now, 22 species have been reported from Brazil from Roraima to Santa Catarina states. Only three additional Brazilian species have been described since Schmid’s (1989) revision: Atopsyche (Atopsaura) blahniki Santos & Holzenthal 2012, Atopsyche (Atopsaura) galharada Santos & Holzenthal 2012 and Atopsyche (Atopsyche) parauna Santos & Holzenthal 2012. The aim of this paper is contribute to the knowledge of Brazilian Atopsyche species, describing three new species from Brazil, adding new distributional notes and providing an identification key for males. The first record of A. rinconi is presented for Brazil, first record of the genus for Midwestern and Northeastern regions of Brazil, as also the records for the states of Bahia, Goiás, and Rio Grande do Sul.

Material and methods Specimens were collected through use of entomological nets, light attraction bed sheets, and Malaise and UV Light Pan traps (Calor & Mariano 2012). The specimens collected by Malaise and UV Light Pan traps were conserved in 80% ethanol. Other specimens were collected using ethyl-acetate kill jars and then pinned. In order to observe and illustrate the characters of genitalia, male abdomens were removed and cleared in a 10% KOH solution and stored permanently in microvials with glycerin. All the observations and illustrations were made under a microscope equipped with a drawing tube. The pencil sketches were scanned for use as templates in Adobe® Illustrator® CS5 and Adobe® Photoshop® CS5. Morphological terminology used in this study follows that of Schmid (1989). Acronyms cited in the text are as follows: AMNH = American Museum of Natural History, New York City, New York, USA; CAS = California Academy of Sciences, San Francisco, California, USA; INHS = Illinois Natural History Survey, Champaign, Illinois, USA; IRSBN = Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium; MCZ = Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA; MZSP = Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil; NMNH = National Museum of Natural History (Smithsonian Institution), Washington, District of Columbia, USA; NMW = Naturhistorisches Museum Wien, Vienna, Austria; UFBA = Museu de Zoologia, Universidade Federal da Bahia, Salvador, Brazil; UMSP = University of Minnesota Insect Collection, Saint Paul, Minnesota, USA; ZSZMH = Zoologische Staatsinstitut und Zoologisches Museum, Hamburg, Germany. Holotypes will be deposited at MZSP, and paratypes at MZSP, UFBA, and UMSP, as indicated below. Additional materials from other collections are cited as below. Bold font in “Distribution” sections indicates new state records of the species.

Family Hydrobiosidae Ulmer 1905 Genus Atopsyche Banks Atopsyche Banks 1905, 17 (Type species: Atopsyche tripunctata Banks 1905, original designation); Ross & King 1952, 177

52 · Zootaxa 4139 (1) © 2016 Magnolia Press

GOMES & CALOR

(revision, biogeography); Schmid 1989, 58 (redescription); Flint et al. 1999, 41 (catalog). Dolochorema Banks 1913, 240 (Type species: Dolochorema irregularis Banks 1913, monobasic), as a distinct genus; Ross 1956, 125 (redescription); Schmid 1989, 60 (redescription, reduced to subgenus of Atopsyche). Harpax Müller 1921, 524 (Type species: no species ever included. Preoccupied by Parkinson 1811 and 3 others); Ulmer 1957, 142 (bibliography, placement, to synonymy with Atopsyche). Ventrarma Navás 1924, 76 (Type species: Ventrarma implexa Navás 1924, original designation); Ross 1947, 127 (to synonymy with Atopsyche); Ross & King 1952, 177 (as subgenus); Ross 1953, 288 (to full synonymy with Atopsyche). Atopsychodes Mosely 1949, 37 (Type species: Atopsychodes cira Mosely 1949, original designation); Ross & King 1952, 177 (to synonymy with Atopsyche). Atopsaura Ross 1953, 292 (Type species: Atopsyche hamata Ross & King 1952, original designation), as a subgenus of Atopsyche.

Sternites. In males, sternite VI with mesoventral spinous lobe and sternite VII with shorter mesoventral spinous lobe. Sexually dimorphic females with similar lobes, but only on sternites V and VI (Schmid 1989). Male genitalia. Segment IX in lateral view with sclerotization long ventrally and laterally, attenutated and mostly membranous dorsally except for internal costa supporting entire anterior circumference, also often with longitudinal internal ventrolateral costae (Schmid 1989). Parapods compressed (laterally flattened) and in horizontal bands, relatively long, wide or narrow, straight or sinuous, simple or complex, spineless, spinose or setose (Schmid 1989). Filipods sometimes absent, emerging mesoventral of preanal appendages, elongate or short (Ross & King 1952). Preanal appendages always present at anterolateral margins of segment X and button-shaped, rounded or elongate, frequently covered by stout setae (Ross & King 1952). Inferior appendages each twosegmented, long, slender, with well developed second article located at apex of first article, often beside apicomesal projection of first article (Schmid 1989). Proctiger large and slightly differentiated, forming dorsal flange (Schmid 1989). Phallic apparatus incomplete, without parameres; phallotheca cylindrical, frequently rounded anteriorly with posterior apex located between parapods, composed of two sclerotized, compressed beaks (“becs de la phallothèque” of Schmid 1989) with different shapes, with membranous endotheca arising between them, this endotheca may be simple or with spines or branches ventrally, laterally or dorsally; aedeagus (“phalicata” of Ross 1956) arising from endotheca, stylet-like, compressed (laterally flattened), arched generally ventrad and then caudad, but with different specific forms; thick tenon from ventrobasal portion of phallotheca articulating with basomesal corners of inferior appendages (Schmid 1989).

Atopsyche species groups with occurrence in Brazil Subgenus Atopsyche A. (A.) tripunctata Group: Inferior appendages simple and primitive, each with second article large and triangular (Schmid 1989, pl. XVI, fig. 1). A. (A.) bolivari Group: Second article of each inferior appendage narrower in middle (Schmid 1989) (in Banks 1924, fig. 36).

Subgenus Atopsaura A. (Atopsaura) longipennis Group: Apicomesal edge of first article of each inferior appendage elongate, forming “tweezers” (chelae) with second article (Fig. 1A) (Schmid 1989).

Atopsyche (Atopsaura) acahuana Schmid 1989 Atopsyche acahuana Schmid 1989, 117 (Type locality: Brazil, ES [Espírito Santo], el. 460 m, 15 km SE Santa Teresa, Faz. Santa Clara, 22.iv.1977, C.M. & Flint, O.S.Jr.; holotype depository: MZSP; male). Atopsyche (Atopsaura) acahuana Schmid; Blahnik et al. 2004, 4 (distribution); Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group in which it seems to be isolated ATOPSYCHE OF BRAZIL

Zootaxa 4139 (1) © 2016 Magnolia Press ·

53

(Schmid 1989). The apical half of the phallotheca is obtusely bulbous and membranous; the first article of each inferior appendage is slightly convex subbasoventrally and subapicodorsally and broader than those of most species, especially at 2/3rds distance from its base, its apicomesal angle is extended in a slender lobe parallel with the second article and nearly the same length (Schmid 1989). Description. Overall body color light brown. Frons and vertex of head with elongate, erect dark and light brown, yellow and whitish setae. Antennal scapes with long brown setae, pedicels and basal flagellomeres yellow. Maxillary and labial palps with long brown and yellow setae. Dorsum of thorax covered by elongate, erect dark and light, brown and yellow setae. Legs yellowish brown. Forewings with shades of dark and light brown, erect setae of veins forming pattern of alternate brown and yellow setae, apex with fringe of light brown and yellow setae. Material examined. Holotype: BRAZIL: Espírito Santo, el. 460 m, 15 km SE Sta.Teresa, Faz. Sta. Clara, 22.iv.1977, C.M. & Flint, O.S.Jr. (pinned, MZSP); Minas Gerais, Rio Caraça, near Santa Barbara, 20°01.371’S, 43°28.750’W, el. 728 m, 9.xi.2001, Holzenthal, R.W., Paprocki, H., Blahnik, R., Amarante, M.; Det. Blahnik, R. 2002. UMSP000080667 to 000080672, 4 males, 2 females (pinned, MZSP); Rio de Janeiro, Encontro dos Rios (Macaé/Bonito) 6 km S Lumiar, 22°23.484’S, 42°18.698’W, el. 600 m, 10.iii.2002, Holzenthal, R.W., Blahnik, R., Paprocki, H. & Prather, A., 5 males, 2 females (pinned, MZSP); same data except Rio das Flores, Macaé de Cima, 10 km SE Mury, el. 1000 m, 09.iii.2002, Holzenthal, R.W, Blahnik, R., Paprocki, H. & Prather, A.; Det. Blahnik, R. 2003, UMSP 000088435 to 000088441, 5 males, 2 females (pinned, MZSP). Distribution. Brazil (Espírito Santo, Minas Gerais, Rio de Janeiro).

Atopsyche (Atopsaura) antisuya Schmid 1989 Atopsyche antisuya Schmid 1989, 117 (Type locality: Brazil, Serra do Cipó, MG [Minas Gerais], Km 116, Rio Brauninha, 16.xii.1974, light, Froehlich, C.G.; holotype depository: MZSP; male). Atopsyche (Atopsaura) antisuya Schmid; Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and can be recognized by the apex of the phallotheca which bears 2 bifid and divergent branches (Schmid 1989). Description. Overall body color yellow to light brown. Frons and vertex of head with elongate, erect light and dark brown, yellow and white setae. Antennal scapes with elongate brown and yellow setae, pedicels and basal flagellomeres yellow, apical flagellomeres brown. First and second segments of maxillary palps with long brown setae. Dorsum of thorax with dark brown, yellow and whitish setae. Forewings light brown with dark brown shade on ventral margins, covered by small brown and yellow setae, erect setae of veins forming irregular pattern of alternate brown and yellowish setae, apices with fringes of alternating patches of light brown and yellow setae. Genitalia of some individuals with filipods slightly longer than preanal appendages. Material examined. BRAZIL: Minas Gerais, Serra do Cipó, Rio Capivara, 06.vii.1974, light, Froehlich, C.G., Shimizu, G.Y. et al., 10 males, 3 females (alcohol, MZSP); same data except Serra do Cipó, Cam. Usina, Rio Capivara, 22.ix.1976, light, Froehlich, C.G., Shimizu, G.Y., 2 males, 1 female (alcohol, MZSP); same data except at light, 18:00–20:00, 18.iv.1975, Froehlich, C.G.; Det. Flint, O.S.Jr. 1977, 13 males, 1 female (alcohol, MZSP); same data except São Gonçalo do Rio Preto, Córrego das Éguas, 18°08.716’S, 43°22.157’W, el. 891 m, 14.x.2000, Paprocki, H., Amarante, M. & Isaac; Det. Blahnik, R. 2001 UMSP000046789 to 000046793, 2 males, 3 females (pinned, MZSP). Distribution. Brazil (Minas Gerais).

Atopsyche (Atopsaura) apurimac Schmid 1989 Atopsyche apurimac Schmid 1989, 118 (Type locality: Brazil, RJ [Rio de Janeiro], el. 1180 m, Km 18, 18 km S. of Teresópolis, 18–19.iv.1977, C.M. & Flint, O.S. Jr.; holotype depository: MZSP; male). Atopsaura (Atopsaura) apurimac Schmid; Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and can be recognized by the long bifid shape of the parapods (Schmid 1989). Description. Overall body color yellowish. Head and thorax with light and dark brown and white setae.


GOMES & CALOR

Antennal scapes yellow with light brown and yellow setae. Pedicels and basal flagellomeres yellow, apical flagellomeres brown. Maxillary and labial palps light brown with light and dark brown setae. Legs yellow. Forewings light brown covered by short light and dark brown setae, light brown and yellowish setae along costal margins and dark brown patch in middle and anal margins of wings, apices with fringes of alternating patches of light and dark brown and yellowish setae. Abdomen anterolaterally with broad glands on terga III and IV; mesolateral small brown setae, and posterolateral long brown setae on terga III, IV, V, VI and VII; lateral margins of proctiger with elongate, brown setae. Material examined. Holotype: BRAZIL: Rio de Janeiro, el. 1180 m, Km. 17, 18 km S of Teresópolis, 1819.iv.1977, C.M. & Flint, O.S. Jr. (pinned, MZSP); Rio Grande do Sul, São José dos Ausentes,15.xii.2007, Melo, A.S. & Ferro, V.G., 1 male (pinned, UFBA). Distribution. Brazil (Rio de Janeiro, Rio Grande do Sul).

Atopsyche (Atopsaura) blahniki Santos & Holzenthal 2012 Atopsyche (Atopsaura) blahniki Santos & Holzenthal 2012, 66 (Type locality: Brazil, Rio de Janeiro, Cachoeiras de Macacu, Rio Souza, 16º26.567'S, 42º37.957'W, el. 150 m, 16.iii.1996, Holzenthal, R., Rochetti & Oliveira; holotype depository: MZSP; male).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and resembles A. zernyi, also described from Brazil. Both species have two long processes in the phallotheca, with the apex upturned and narrow (Santos & Holzenthal 2012). The apex of the phallotheca has long, paired processes with several spines on their lateral edges (Santos & Holzenthal 2012). Description. See Santos & Holzenthal (2012). Material examined. None. Distribution. Brazil (Rio de Janeiro).

Atopsyche (Atopsyche) chirihuana Schmid 1989 Atopsyche chirihuana Schmid 1989, 121 (Type locality: Ecuador, Pich., Santo Domingo (47 km S), Rio Palenque Bio. Station, el. 750 m; holotype depository: NMNH; male). Atopsyche (Atopsyche) chirihuana Schmid; Blahnik et al. 2004, 4 (distribution); Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsyche) tripunctata Group and is most similar to A. erigia Ross 1947. It can be recognized by the angled second article of each inferior appendage (Schmid 1989). Description. Overall color yellowish brown; frons and vertex of head with elongate, erect light and dark brown, yellow and whitish setae. Antennal scapes with long brown setae. Pedicels and basal flagellomeres yellow, apical flagellomeres brown. Dorsum of thorax with elongate, erect light brown and whitish setae. Forewings light brown with brown shading on ventral margin, covered by small light brown and whitish setae, erect setae of veins forming irregular pattern of alternate brown and whitish setae, apices with fringes of light brown and yellow setae; legs yellow to light brown. Genitalia, inferior appendages large and setose in anterior region, each with second article about half-length of first and obtusely curved apically (Schmid 1989). Material examined. BRAZIL: Minas Gerais, Rio Santo Antonio, downstream from Morro do Pilar, 19°08.134’S, 43°21.256’W, el. 530 m, 17.x.2000, Paprocki, H. & Ferreira; Det. Blahnik, R. 2001, UMSP 000047568, 1 male (pinned, MZSP). Distribution. Brazil (Minas Gerais), Ecuador.

Atopsyche (Atopsyche) erigia Ross 1947 Atopsyche erigia Ross 1947, 129 (Type locality: Mexico, Edo. Tamaulipas, Hacienda Santa Engracia, 09.iii.1939; holotype depository: INHS; male); Edwards & Arnold 1961, 401 (larva; distribution); Bueno-Soria & Flint 1980, 191 (distribution). Atopsyche (Atopsyche) erigia Ross; Blahnik et al. 2004, 4 (distribution); Paprocki et al. 2004, 7 (checklist); Calor 2011, 320 (checklist). ATOPSYCHE OF BRAZIL


55

Diagnosis. This species belongs to the Atopsyche (Atopsyche) tripunctata Group and is recognized by the parapods each having a short subapicodorsal tooth and another short apicodorsal tooth; the filipods being longer than the parapods; the first article of each inferior appendage being straight, parallel-sided and simple; and the phallotheca being broad and sinuate, truncate apically (Ross 1947). This species is most similar to A. chirihuana in having parapods each with two short dorsal teeth, a simple phallotheca, and a subtriangular second article of each inferior appendage; however, in A. erigia the second article is rounded apically, not truncate as for A. chirihuana. Description. Overall body color yellowish to dark brown. Antennal scapes with light brown and white setae, pedicels and basal flagellomeres yellow, apical flagellomeres brown. Maxillary and labial palps with light brown and yellow setae. Setae of head and thorax elongate, erect, whitish with few darker setae. Postero- and mesolateral elongate setae in all abdominal terga. Legs yellowish to dark brown. Forewings light brown with yellowish setae at midlength, apices with fringes of alternating patches of brown and and yellowish setae; posterolateral glands on tergum II. Genitalia, with long slender filipods bearing scattering of long setae at apices, parapods upcurved and each bearing dorsal indentation and then tooth some distance before apex, inferior appendages long and slender, each with first article more than twice as long as wide; second article evenly tapering and rounded at apex, little more than one-half as long as first article; phallic apparatus broad and sinuate, truncate at apex, aedeagus upturned and curved (Ross 1947). Material examined. BRAZIL: Minas Gerais, Serra do Cipó, Rio Cipó in Cardeal Mota, (Cachoeira baixo), 19°20.553’S, 43°38.531’W, el. 750 m, 10.xi.2001, Holzenthal, R., Paprocki, H., Blahnik, R., Amarante, M.; Det. Blahnik, R., 2002, 00079869 to 000079902, 8 males, 10 females (pinned, MZSP); São Paulo, Altinópolis, Fazenda São João da Mata, Rio Baguassu, 21°00.588’S, 47°28.900’W, el. 745 m, 19-21.xi.2003, Holzenthal, R.W., Paprocki, H. & Calor, A.; Det. Blahnik, R., 2007, UMSP 000119778 to 0001197941, 7 males (pinned, MZSP); same data except 000120546 to 000120579, 5 males, 10 females (pinned, MZSP). Distribution. Brazil (Minas Gerais, São Paulo), Costa Rica, Guatemala, Mexico, Nicaragua, Panama, USA.

Atopsyche (Atopsaura) galharada Santos & Holzenthal 2012 Atopsyche (Atopsaura) galharada Santos & Holzenthal 2012, 71 (Type locality: Brazil, São Paulo, Campos do Jordão, Parque Estadual de Campos do Jordão, Rio Galharada, 22º41.662’S, 45º27.783’W, el. 1530 m; holotype depository: MZSP; male).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group. It resembles A. ayahuaca Schmid 1989 and A. plancki Marlier 1964 in the absence of filipods and in the simple phallotheca (Santos & Holzenthal 2012). Parapods each have two carinae bearing tooth-like processes and the apicomesal process of the first article of each inferior appendage is very long. Moreover, A. galharada has a broad lateral sclerotized projection on each side of the proctiger and the phallotheca has two long processes (“beaks” of Schmid 1989) broadest subapicolaterally, in dorsal view (Santos & Holzenthal 2012). Description. See Santos & Holzenthal (2012). Material examined. BRAZIL: São Paulo, Campos do Jordão, P. E. Campos do Jordão, Córrego Casquilho, 20.xi.2007, light, Pinho, L., Mariano, R. & Lecci, L, 1 male (alcohol, UFBA); same data except Afluente Córrego Garalhada, 22º41’33”S 45º27’55”W, 05.v.2001, U.V./White light, Calor, A.R., Lecci, L. & Moretto, R., 1 male (pinned, UFBA). Distribution. Brazil (São Paulo).

Atopsyche (Atopsaura) hamata Ross & King 1952 Atopsyche hamata Ross & King 1952, 202 (Type locality: Brazil, Summit of Mt. Roraima; holotype depository: AMNH; male). Atopsyche (Atopsaura) hamata Ross & King; Ross 1953, 292, type species of subgenus Atopsyche (Atopsaura); Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group. It is closely similar to A. longipennis in that the filipods are about as long as the parapods and the apicomesal portion of the first article of


GOMES & CALOR

each inferior appendage is about as long as the second article of the same appendage and completely overlapped by it (Ross & King 1952). Atopsyche hamata also resembles A. antisuya in the apical portion of the first article and in the second article of an inferior appendage, but differs in that the first article is curved caudoventrad in A. hamata. Description. Parapods simple, each widest at center, a single small dorsal tooth near apex. Filipods sinuate, about as long as parapods. Inferior appendages each with first article cylindrical, curved and mesal apex prolonged into fingerlike tip; second article as long as, and completely overlapping prolongation of first. Principal lobes (“beaks” of Schmid 1989) of phallic apparatus each bearing short, sclerotized spine on its posteroventral angle and long, petiolate membranous process (endotheca) on its posterodorsal angle, this process more than 1/3rd as long as entire phallotheca, bearing short spine basoventrally resembling and perpendicular to those borne ventrally on principal lobes and smaller spines and setae on apical third of process; aedeagus L-shaped (Ross & King 1952). Material examined. None. Distribution. Brazil (Roraima).

Atopsyche (Atopsaura) hatunpuna Schmid 1989 Atopsyche hatunpuna Schmid 1989, 122 (Type locality: Brazil, Casa Grande, S.P. [São Paulo], Ribeira, Coruja; holotype depository: MZSP; male). Atopsyche (Atopsaura) hatunpuna Schmid; Paprocki et al. 2004, 7 (checklist); Dumas et al. 2010, 7 (distribution); Calor 2011, 320 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group. It resembles A. huarcu Schmid 1989 and A. huamachucu Schmid 1989 in the apical portions of the phallotheca forming two concave processes (“beaks”), each with an internal carina (Schmid 1989), and a dorsal process. Atopsyche hatunpuna can be recognized by the shape of its dorsal process, which is compressed (laterally flattened) and broadened in it apical half with an acute apex (Schmid 1989). Description. See Schmid (1989). Material examined. None. Distribution. Brazil (Minas Gerais, São Paulo).

Atopsyche (Atopsaura) huacachaca Schmid 1989 Atopsyche huacachaca Schmid 1989, 122 (Type locality: Brazil, Rio de Janeiro, Itatiaia, el. 1700 m, Registro Pass, 18.x.1985, Miller, S.E.; holotype depository: MZSP; male). Atopsyche (Atopsaura) huacachaca Schmid; Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and is most similar to A. sanctipauli and A. huarcu. This similarity is based on the rounded anterodorsal portion of the second article of each inferior appendage and on the similarly shaped dorsal process of the phallotheca (Schmid 1989). The parapods form simple obtuse triangles in lateral view with apical setae and each with a highly obtuse anterodorsal angle and a dozen, small, thick tooth-like spines on the dorsal edge (Schmid 1989). Description. Overall body color yellowish to light brown. Head and thorax covered by elongate, erect light and dark brown and whitish setae. Antennal scapes light brown with elongate dark brown setae. Pedicels yellow, basal flagellomeres light brown and apical flagellomeres brown; maxillary and labial palps light brown with long light and dark brown setae. Legs yellowish to light brown. Forewings light brown, erect setae of veins forming irregular pattern of alternate black, dark and light brown and yellow setae, apices with fringes of alternating patches of light and dark brown. Material examined. Holotype. BRAZIL: Rio de Janeiro, Itatiaia, el. 1700 m, Registro Pass., 18.x.1985, Miller, S.E. (pinned, MZSP). Distribution. Brazil (Rio de Janeiro).

ATOPSYCHE OF BRAZIL


57

Atopsyche (Atopsaura) huamachucu Schmid 1989 Atopsyche huamachucu Schmid 1989, 124 (Type locality: Brazil, RJ [Rio de Janeiro], el. 1180 m, km 17, 18 km S of Teresópolis; holotype depository: MZSP; male). Atopsyche (Atopsaura) huamachucu Schmid; Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and is similar to A. sanctipauli and A. huachacuyac Schmid 1989. The phallotheca has a membranous anterodorsal portion; in lateral view, each “beak” appears as a simple lobe with a narrowly oval apex, having an internal carina with a minuscule preapical point in dorsal view; moreover a basodorsal process of the phallotheca is very regularly slender, arched basally upward and then downward, its slightly variable length surpassing the apices of the inferior appendages (Schmid 1989). Description. Overall body color yellowish to light brown. Antennal scapes light brown with elongate light brown setae, pedicels and basal flagellomeres light brown. Setae of head and thorax whitish with some darker setae. Maxillary and labial palps light brown with long dark brown setae; forewings brown and irregular shades of brown, covered by small brown and yellow setae, erect dark and light brown and yellow setae on veins, apices with fringes of dark brown and yellowish setae. Legs yellowish to light brown. Material examined. Holotype. BRAZIL: Rio de Janeiro, Km. 17, 18 Km S. of Teresópolis, 18-19.iv.1977, Froehlich, C.G. & Flint, O.S. Jr. (pinned, MZSP). Distribution. Brazil (Rio de Janeiro).

Atopsyche (Atopsaura) huanapu Schmid 1989 Atopsyche huanapu Schmid 1989, 124 (Type locality: Brazil, São Paulo, Salesópolis, Estação Biológica de Boracéia, Parede da Pedreira; holotype depository: MZSP; male). Atopsyche (Atopsaura) huanapu Schmid; Blahnik et al. 2004, 4 (distribution); Paprocki et al. 2004, 7 (checklist); Calor 2011, 320 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and is similar to A. serica and A. muelleri n. sp. in the concave mesal surface of the second article of each inferior appendage and the upcurved apicodorsal portion of each parapod (Schmid 1989). It is characterized by the slender shape of the apex of the phallotheca in lateral view and its V shape in dorsal view and by the distinctive shapes of the first article apex and the second article of each inferior appendage (Schmid 1989). Description. Overall body color light to dark brown. Head and thorax with light and dark brown and yellow elongate, erect setae. Maxillary and labial palps light brown with dark brown setae; antennal scapes dark brown with brown setae, pedicels light brown, basal flagellomeres yellow and distal flagellomeres brown. Legs light brown, coxae of all legs and fore- and midleg tibiae darker brown. Forewings brown covered with light and dark setae, apices with fringes of light brown setae. Material examined. Holotype. BRAZIL: São Paulo, Estação Biológica de Boracéia, Parede da Pedreira, 30.ix.1966, Froehlich, C.G. (alcohol, MZSP). Distribution. Brazil (Rio de Janeiro, São Paulo).

Atopsyche (Atopsaura) huarcu Schmid 1989 Atopsyche huarcu Schmid 1989, 125 (Type locality: Brazil, Minas Gerais, Nova Lima, el. 850 m; holotype depository: MZSP; male). Atopsyche (Atopsaura) huarcu Schmid; Blahnik et al. 2004, 4 (distribution); Paprocki et al. 2004, 7 (checklist); Calor 2011, 320 (checklist).

Diagnosis. This species is classified in the Atopsyche (Atopsaura) longipennis Group and is similar to Atopsyche kamakan n. sp. and A. sanctipauli in the basomesal concavity of the second article of each inferior appendage and in the presence of a dorsal process of the phallotheca; however the apical portion of the first article of the inferior


GOMES & CALOR

appendage is broader dorsally in A. huarcu and the anterior dorsal tooth on each parapod is much larger and more nearly anterior. The dorsal process of the phallotheca is enlarged apically in A. sanctipauli and A. huarcu and all three species have one or more teeth on the dorsal edge of this process, but the teeth are small and evenly distributed over about half of the length of the process in A. huarcu, whereas there is only a single tooth on the A. kamakan process and the teeth on the process of A. sanctipauli are large and arranged in a fan-like array. Description. Overall body color yellowish brown; legs yellow to light brown. Head and thorax with long, erect light brown, yellow and whitish setae. Antennal scapes yellow with long light brown setae. Maxillary and labial palps with light brown setae. Forewings light brown, covered by light brown setae, erect setae of veins forming irregular pattern of alternate light and dark brown setae. Broad anterolateral glands on tergum IV; long brown lateral and posterolateral setae on terga IV, V, VI, VII and VIII; proctiger with long brown setae. Material examined. Holotype. BRAZIL: Minas Gerais, Nova Lima, el. 850 m, 21.x.1985, Miller, S.E. (pinned, MZSP). Distribution. Brazil (Minas Gerais, Rio de Janeiro, São Paulo).

Atopsyche (Atopsaura) longipennis (Ulmer 1905) Psilochorema longipennis Ulmer 1905, 110 (Type locality: not given; type depository: ZSZMH; male). Atopsyche brasiliana; Ulmer 1909, 73 [a lapsus for Atopsyche longipennis (Ulmer)]. Atopsyche longipennis (Ulmer); Ross & King 1952, 202 (type from Brazil, Santa Catarina; male); Flint 1972, 225 (distribution). Atopsyche (Atopsaura) longipennis (Ulmer); Paprocki et al. 2004, 7 (checklist).

Diagnosis. Atopsyche longipennis belongs in the Atopsyche (Atopsaura) longipennis Group and closely resembles A. hamata in that the filipods are about as long as the parapods, and the apicomesal projection of the first article of each inferior appendage is similar in shape (Ross & King 1952). This species can be recognized by the boot-shaped second article of each inferior appendage (Ross & King 1952). Description. Overall body color yellowish to light brown. Setae of head and thorax light to dark brown with long light and dark brown setae. Antennal scapes light brown with dark brown and yellow setae, pedicels and basal flagellomeres yellow and apical flagellomeres light brown. Legs yellow to light brown. Forewings light brown with yellowish setae along costal margin and light brown erect setae on ventral margin. Material examined. BRAZIL: Santa Catarina, Morro do Baú, Mun. Ilhota, 03.xii.1975; Det. Flint, O.S. Jr., 1977, 1 male (alcohol, MZSP). Distribution. Argentina, Brazil (Santa Catarina).

Atopsyche (Atopsyche) parauna Santos & Holzenthal 2012 Atopsyche (Atopsyche) parauna Santos & Holzenthal 2012, 69 (Type locality: Brazil, Minas Gerais, Rio Paraúna, 3 km Santana do Riacho, 19º10.986’S, 43º43.485’W, el. 650 m; holotype depository: MZSP; male).

Diagnosis. This species belongs to the Atopsyche (Atopsyche) tripunctata Group. It resembles A. jaba Blahnik & Gottschalk 1997, described from Costa Rica, in the following characters: The apicomesal processes of the first article of each inferior appendage are similar in shape and the “beaks” of the phallotheca are each divided into two long, paired processes posteriorly (Santos & Holzenthal 2012). Atopsyche parauna resembles A. urumarca Schmid 1989 in the second article of each inferior appendage being slightly arched; however this species can be distinguished from A. urumarca by the divided shape of the “beaks” of A. parauna. This species can be recognized also by the broad parapods, each with a serrate dorsolateral margin and a midlateral spinous projection; the short and slightly hooked second article of each inferior appendage; and the phallotheca is posteriorly divided into two long paired processes, with the dorsal one bifurcate apically and the ventral one curved mesad, dorsal view (Santos & Holzenthal 2012). Description. See Santos & Holzenthal (2012). Material examined. None. Distribution. Brazil (Minas Gerais). ATOPSYCHE OF BRAZIL


59

Atopsyche (Atopsaura) plancki Marlier 1964 Atopsyche plancki Marlier 1964, 2 (Type locality: Brazil, São Paulo, Salesópolis, Estação Ecológica de Boracéia; holotype depository: IRSNB; male). Atopsyche (Atopsaura) plancki Marlier; Blahnik et al. 2004, 4 (distribution); Paprocki et al. 2004, 7 (checklist); Santos & Holzenthal 2012, 77 (distribution); Calor 2011, 320 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group (Schmid 1989) and resembles A. galharada in the short and broad first article of each inferior appendage. Atopsyche plancki can be recognized from its congeners by the first article of each inferior appendage short and broad, with an acute apicoventral process. Description. Overall body color yellow to light brown. Frons and vertex of head with elongate, erect, brown and whitish setae; dorsum of thorax covered by elongate, erect brown and whitish setae. Antennal scapes with long light and dark brown and yellowish setae, pedicels and basal flagellomeres yellow and apical flagellomeres brown. Maxillary and labial palps with long, brown setae. Forewings light brown with darker shade of brown in posteroventral margin, erect setae of veins forming irregular pattern of alternate brown and yellow setae, apex with fringe of alternating patches of light brown and yellow setae; proctiger, in lateral view, very broad apically with dorsal and posteroventral margin covered with long brown setae. Material examined. BRAZIL: Minas Gerais, Rio Caraça, near Santa Barbara, 20°01.371’S, 43°28.750’W, el. 728 m, 09.xi.2001, Holzenthal, R., Paprocki, H. Blahnik, R. & Amarante, M.; Det. Blahnik, R., 2002, UMSP 000080620 to 000080637, 5 males, 13 females (pinned, MZSP); Rio de Janeiro, Rio Macaé, Macaé de Cima, 22°23.684’S, 42°30.136’W, 1000 m, 08.iii.2002, Holzenthal R., Blahnik, R., Paprocki, H., Prather, A.; Det. Blahnik, R., 2003 UMSP 000086049 e 000086050, 2 males (pinned, MZSP); same data except Rio das Flores, Macaé de Cima, 10 Km SE Mury, el. 1000 m, 09.iii.2002, Holzenthal R., Blahnik, R., Paprocki, H. & Prather, A.; Det. Blahnik, R., 2003, UMSP 000088420 to 000088424, 1 male, 4 females (pinned, MZSP); Santa Catarina, Parque Ecológico Spitzkopf, confl. Rio Ouro & Rio Caeté 27°00.352’S, 49°06.693’W, el. 140 m, 25.xi.2003, Holzenthal R., Paprocki, H. & Calor, A.R., 3 males (alcohol, MZSP); São Paulo, Iporanga, Parque Estadual Intervales, 16.xii.1999, light, Bispo, P. & Crisci-Bispo, V., 1 male (alcohol, MZSP). Distribution. Brazil (Rio de Janeiro, Santa Catarina, São Paulo).

Atopsyche (Atopsyche) rinconi Holzenthal & Cressa 2002 Atopsyche (Atopsyche) rinconi Holzenthal & Cressa 2002, 138 (Type locality: Venezuela, Rio Cocolllar [sic]; 1.5 km SE Las Piedras de Cocollar, 10°09.617”N, 63°47.605”W, el. 810 m, 7–8.iv.1995, Holzenthal & Flint; holotype depository: NMNH; male).

Diagnosis. This species belongs in the Atopsyche (Atopsyche) tripunctata Group and is most similar to A. chirihuana and A. erigia, especially in the wide second article of each inferior appendage, the long filipods, and the broad phallotheca in lateral view. Segment IX is most similar to that of A. chirihuana by the sinuous anterolateral margin, however segment IX is slightly shorter longitudinally. Dorsal processes on the parapods are upcurved. The first article of each inferior appendage is short, and the apex is broad and rounded. Description. Overall body color yellow to brown. Head, frons and vertex with brown setae. Antennal scapes light brown with small brown setae, pedicels yellow, basal flagellomeres yellow, apical flagellomeres brown. Maxillary palps absent (broken off); labial palps brown with long brown and yellow setae. Thorax with light and dark brown setae. Legs yellowish brown, midcoxae and foreleg tibiae darker brown, mid- and hind leg tibiae with yellow setae. Forewings light brown, with shades of darker brown, especially on Sc vein, erect light brown and yellow setae. Terga III to VIII with elongate brown setae in lateral margins; tergum III with pair of prominent, rounded glands located at anterolateral margins; sterna II to VII with numerous minute setae; sternum VIII with long brown setae; sternum VI ventromesal process long, slender, curved, with short spine-like setae along ventral margin and terminating with longer, spine-like setae; ventromesal process on sternum VII shorter than anterior process, straight. Material examined. BRAZIL: Bahia, Abaíra, Estrada Velha Ouro Verde-Piatã, Rio Toboro, 28.vii.2010, UV Light Pan trap , Calor, A.R., Quinteiro, F.B., França D., Lecci, L.S., Arantes, T. & Camelier, P., 3 males (alcohol,


GOMES & CALOR

UFBA); Wenceslau Guimarães, Estação Ecológica de Wenceslau Guimarães, Riacho Serra Grande, 10.x.2010, UV Light Pan trap, Calor, A.R., Quinteiro, F.B. & Costa, A.M., 1 male (alcohol, UFBA). Minas Gerais, Cabo Verde, Pedregal, 21°28’06”S, 46°24’13”W, el. 920 m, 02–05.xi.2006, light, Amorim, D., Falaschi, R. & Oliveira, S., 27 males (alcohol, UFBA); same data except, 2 males (pinned, UFBA). Distribution. Venezuela, Brazil (Bahia, Minas Gerais).

Atopsyche (Atopsaura) sanctipauli Flint 1974 Figures 1A–B Atopsyche sanctipauli Flint 1974, 5 (Type locality: Brazil, São Paulo, Alto da Serra; holotype depository: NMW; male). Atopsyche (Atopsaura) sanctipauli Flint; Blahnik et al. 2004, 4 (distribution); Paprocki et al. 2004, 7 (checklist); Calor 2011, 320 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and is most similar to A. japoda Ross & King 1952 in the dorsal process on the phallotheca. This new species also is similar to Atopsyche kamakan n. sp. in and A. huarcu in the basomesal concavity of the second article of each inferior appendage and in the presence of a dorsal process of the phallotheca. However, it can be recognized by the thin forked lateral process of the phallotheca connected laterally, by the small lobe from the mesal face of each inferior appendage and by the shapes of the apical pieces of the inferior appendages (Flint 1974). Description. Atopsyche sanctipauli shows variations in the shape of the parapods, but generally this structure is broad basally, each with a basodorsal process directed anterad, tapering to a narrow posterior apex bearing setae, dorsal margin with spines (Fig. 1A). Phallotheca with nonarticulated dorsal process; “beaks” thin, each with narrow dorsolateral flange (Flint 1974, fig. 27), laterally with thin, deeply bifurcate process dorsally with long, slender, central spine (aedeagus in Schmid’s nomenclature); aedeagus nearly as long as phallotheca, slender, curved ventrad and then caudad, apically acute (Fig. 1B; Flint 1974). The dorsal central spine shows morphological variation in some specimens which have a conspicuous dorsal array of sharp spines near its middle. Material examined. BRAZIL: Minas Gerais, Serra do Cipó, Cardeal Mota, 19°18.912’S, 43°36.260’W, el. 800 m, 12.xi.2001, Holzenthal, R.W., Paprocki, H., Blahnik, R. & Amarante, M.; Det. Blahnik, R., 2002, UMSP 000081910 to 000081931, 16 males, 6 females (pinned, MZSP); same data except Rio Capivara, light, 18.xii.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 1 male (alcohol, MZSP); same data except Km 126, 17.iv.1975, light, Froehlich, C.G., Carvalho, C., Shimizu, G.; Det. Flint, O.S. Jr., 1 male (alcohol, MZSP); same data except light, 17.xii.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 2 males (alcohol, MZSP); same data except afl. Rio Capivara, 19.iv.1975, light, Froehlich, C.G., Carvalho, C., Shimizu, G.; Det. Flint, O.S. Jr., 1977, 1 male (alcohol, MZSP); Rio de Janeiro, Parati, trib. to Riacho Perequê-açu, 23°12.833’S, 44°47.483’W, el. 190 m, 26.ix.2002. Blahnik, R., Prather, A., Melo, A., Froehlich, C.G. & Silva, R.; Det. Blahnik, R., UMSP 000087680, 1 male (pinned, MZSP); Rio Grande do Sul, São José dos Ausentes, 28°57’659”S, 49°87’795”W, el. 1178 m, 21.xi.2007, Melo, A. & Ferro, V., 1 male (pinned, UFBA); same data except 28°57’319”S, 49°84’292”W, el. 1250 m, 08.ii.2008, 1 male (pinned, UFBA); same data except 09.ii.2008 28°57’607”S, 49°92’000”W, el. 1250 m, 1 male (pinned, UFBA); same data except 11.xi.2007, 2 males (pinned, UFBA); same data except 28°57’319”S, 49°84’292”W, el. 1250 m, 08.iii.2008, 4 males (pinned, UFBA); same data except 28°60’407”S, 49°76’236”W, el. 1238 m, 09.iii.2008, 5 males (pinned, UFBA); Santa Catarina, Florianópolis, Rio Estaleiro, 27°47’49”S, 48°33’28”W, 06–12.x.2012, Malaise trap, Parise, A., 1 male (alcohol, UFBA); same data except 12-21. X.2012, 1 male (alcohol, UFBA); same data except 27°36’35”S, 48°30’57”W, 27.ix–24.x.2012, Pinho, L. et al., 1 male (alcohol, UFBA); Grão Pará, P.E. Serra Furada, #01, 28°11’28”S, 49°23’30”W, 13.x.2012, light trap, Pinho, L., Ganzer, A., Gomes, L. & Parise, A., 1 male (alcohol, UFBA); Urubici, Cascata Avencal #30, 28°03’00”S, 49°37’02”W, 09.i.2013, light trap, Pinho, L., Novaes, M. & Haddad, M., 3 males (alcohol, UFBA); same data except Rio Canoas, #29, 28°01’41”S, 49°22’36”W, 08.i.2013, Pinho, L., Novaes, M. & Haddad, M., 1 male (UFBA); São Paulo, Estação Biológica Boracéia, Rio Venerando, 23°39.185’S, 45°53.414’W, el. 850 m, 18– 21.ix.2002, Blahnik, R., Prather, A., Melo, A., Froehlich, C.G. & Silva, R.; Det. Blahnik, R., 2003, UMSP 000088080, 1 male (pinned, MZSP); Casa Grande, Pedreira, 15.xi.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 2 males, 1 female, (alcohol, MZSP); same data except Rib. Coruja, light, 26.i.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 3 males, 1 female (alcohol, MZSP); same except except light, 12.x.1974; Det. Flint, O.S. Jr., 1977, 1 ATOPSYCHE OF BRAZIL


61

male, 7 females (alcohol, MZSP); same data except 16.xi.1974, Froehlich, C.G; Det. Flint, O.S. Jr., 1977, 2 males, 15 females (alcohol, MZSP); same data except 03.xi.1969, light; Det. Flint, O.S. Jr., 1977, 1 male, 2 females (alcohol, MZSP); same data except Casa Grande, Pedreira, light, 13.x.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 1 male, 6 females (alcohol, MZSP); same data except Iporanga, Parque Estadual Intervales, 16.xii.1999, light, Bispo, P. & Crisci-Bispo, V., 1 male (alcohol, MZSP); same data except 08.i.2000, light, Bispo, P., CrisciBispo, V., 1 male (alcohol, MZSP); same data except 17.vii.1999, light, Bispo, P., Crisci-Bispo, V., 1 male (alcohol, MZSP); same data except 15.vii.1999, light, Bispo, P., Crisci-Bispo, V., 8 males (alcohol, MZSP). Distribution. Brazil (Minas Gerais, Paraná; Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo).

FIGURE 1. Atopsyche (Atopsaura) sanctipauli Flint, male. 1A, genitalia, left lateral, with inset of apex of right inferior appendage, mesal; 1B, phallic apparatus, left lateral. 1st article = first article of an inferior appendage; 2nd article = second article of an inferior appendage; ae = aedeagus; beaks = beaks of the phallotheca; d proc = dorsal process of the phallotheca; IX = segment IX; lobe = lobe of the phallotheca; par = left parapod; phal = phallic apparatus; pr app = left preanal appendage; proc = proctiger.

Atopsyche (Atopsaura) serica Ross 1953 Atopsyche (Atopsaura) serica Ross 1953, 292 (Type locality: Brazil, Nova Teutônia, 27°11’S, 52°23’W, 04.x.1939, Fritz Plaumann; holotype depository: MCZ; male); Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and most closely resembles A. longipennis in the short parapods and filipods and A. hamata in the curved first article of each inferior appendage. It is distinguished from them by an extra pair of short lateral processes below the filipods and a sclerotized horseshoe-shaped or hood-like structure above the opening for the phallic apparatus (Ross 1953). Description. Frons and vertex of head with elongate, erect brown and whitish setae; dorsum of thorax covered by elongate, erect, brown, whitish and yellowish setae; forewings brown, covered by short brown and yellow setae,


GOMES & CALOR

erect setae of veins forming irregular pattern of alternating brown and yellow setae, apices with fringes of alternating patches of brown and yellow setae. Material examined. BRAZIL: Rio Grande do Sul, São José dos Ausentes, 8°57’6.07”S, 49°92’0.00”W, el. 1250 m, 11.xii.2007, Melo, A. & Ferro, V., 3 males (pinned, MZSP); same data except 28.61973S, 49.85281W, el. 1205 m, 19.i.2007, Melo, A. & Ferro, V., 3 males (pinned, MZSP); Santa Catarina, Parque Ecológico Spitzkopf, confl. Rio Ouro & Rio Caeté 27°00.352’S, 49°06.693W, el. 140 m, 25.xi.2003, Holzenthal, R.W., Paprocki, H. & Calor, A.R., 4 males, 1 female (pinned; MZSP); Blumenau, 04.xii.1975, Exc. Dep. Zool. USP, 2 males, 2 females (alcohol, MZSP). Distribution. Brazil (Rio Grande do Sul, Santa Catarina).

Atopsyche (Atopsaura) siolii Flint 1971 Atopsyche siolii Flint 1971, 12 (Type locality: Brazil, Rio Marauiá, Cachoeira Pora Comeschie, dichtoberhalb Endstation, Benthos aus der Strömung, 28.i.1963, E.J. Fittkau; holotype depository: NMNH; male, larva, pupa). Atopsyche (Atopsaura) siolii Flint; Paprocki et al. 2004, 7 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and resembles A. muelleri n. sp. in the similarly shaped apicodorsal process of each parapod, the short filipods and the forked apical portion of the phallotheca. Atopsyche siolii differs from other species in the genus by the very broad and rather short parapods, the shape of the “beaks” of the phallotheca, and the broader second article of each inferior appendage (Flint 1971). Description. See Flint (1971). Material examined. None. Distribution. Brazil (Amazonas).

Atopsyche (Atopsyche) urumarca Schmid 1989 Atopsyche urumarca Schmid 1989, 131 (Type locality: Brésil [Brazil], Serra do Cipó, M.G. [Minas Gerais], Rio Capivara, 08.vii.1974, Luz, Froehlich, C.G. et al.; holotype depository: MZSP; male). Atopsyche (Atopsyche) urumarca Schmid; Paprocki et al. 2004, 7 (checklist); Santos & Holzenthal 2012, 75 (distribution; variation; male).

Diagnosis. This species belongs to the Atopsyche (Atopsyche) bolivari Group and resembles A. parauna in the small, arched second article of each inferior appendage. Distinctively, the parapods are each broad basally with a variably large or small oblique U-shaped incision at mid length of the dorsal margin in all specimens. The phallotheca is broadly rounded basally and with three paired processes posteriorly: two short, lateral, and one long, dorsal (“beaks”). Each “beak” bearing two small acute projections at the apex, which is truncate to shallowly excised and with the dorsal portion extended in a curved hook (Schmid 1989; Santos & Holzenthal 2012). Description. Overall body color yellow to brown (in alcohol). Setae of head and thorax elongate, erect light and dark brown; antennal scapes brown with long dark brown setae, pedicels and basal flagellomeres yellow and apical flagellomeres light brown. Maxillary and labial palps brown with long light and dark brown setae. Legs yellow to light brown. Forewing light brown with anterior and anal region dark brown, covered by short light and dark brown setae. Remarks. In the genitalia, variation has been observed among populations in southeastern Brazil, especially in the shapes of the parapods and the beaks of the phallotheca (Santos & Holzenthal 2012). Material examined. Holotype. BRAZIL: Minas Gerais, Serra do Cipó, Rio Capivara, 08.vii.1974, light, Froehlich, C.G. et al. (alcohol, MZSP); Paratypes: Minas Gerais, Serra do Cipó, Rio Capivara, 18.xii.1973, light, Froehlich, C.G., 1 male, (alcohol, MZSP); Goiás, Chapada dos Veadeiros, Solário, Moinho, 14.xii.2006, light, Bispo, P.; Yokoyama, E. & Paciencia, G., 3 males (alcohol, UFBA). Distribution. Brazil (Goiás, Minas Gerais, São Paulo).

ATOPSYCHE OF BRAZIL


63

Atopsyche (Atopsaura) usingeri Denning 1968 Atopsyche usingeri Denning 1968, in Denning & Sykora 1968, 172 (Type locality: Brazil, Rio de Janeiro, Teresópolis, Organ Mountains, 30.viii.1957, Usinger, R.; holotype depository: CAS). Atopsyche (Atopsaura) usingeri Denning; Paprocki et al. 2004, 7 (checklist, species credited to Denning & Sykora); Calor 2011, 320 (checklist, species credited to Denning & Sykora).

Diagnosis. This species belongs in the Atopsyche (Atopsaura) longipennis Group. It resembles A. serica and A. huanapu in the short parapods which in lateral view are parallel-sided, each with its apex truncate and its apicodorsal angle subacute and upturned. It differs from A. serica in that the filipods of A. usingeri are much longer than the parapods, slender, slightly enlarged in their apical portion. Atopsyche usingeri differs from both of these species by lacking a transverse sclerite above the phallotheca (Schmid 1989), in the different horseshoe-shaped or hood-like structure of A. serica (Ross 1953), and in the dorsal process on the phallotheca. Description. Overall body color light to dark brown (alcohol). Head with light and dark brown and whitish setae. Antennal scapes brown with brown setae, pedicels and basal flagellomeres light brown, apical flagellomeres brown. Maxillary and labial palps light brown with elongate dark brown setae. Dorsum of thorax with dark and light brown setae. Forelegs and hind legs dark brown, midleg light brown. Forewings light brown, covered with small light and dark brown setae (in alcohol). Terga II to VIII with dorsal and lateral long brown setae. Remarks. Some individuals have the apex of the first article of each inferior appendage slightly broader than the basal portion, differring from the illustration and description of Denning & Sykora (1968), and the second article is sometimes downcurved. Material examined. BRAZIL: São Paulo, E.B. Paranapiacaba, 17.ix.1963; Det. Flint, O.S. Jr., 1977, 2 males (alcohol, MZSP); same data except 27.viii.1963, light; Det. Flint, O.S. Jr., 1977, 1 male (alcohol, MZSP); same data except 27.xi.1963, light; Det. Flint, O.S. Jr., 1977, 1 male (alcohol, MZSP); same data except 05.vii.1963, light; Det. Flint, O.S. Jr., 1977, 1 male (alcohol, MZSP). Distribution. Brazil (Rio de Janeiro, São Paulo).

Atopsyche (Atopsaura) zernyi Flint 1974 Figures 2A–H Atopsyche zernyi Flint 1974, 5 (Type locality: Brazil, São Paulo, Alto da Serra, 29-30.x.1927; holotype depository: NMW; male). Atopsyche (Atopsaura) zernyi Flint; Blahnik et al. 2004, 4 (distribution); Paprocki et al. 2004, 7 (checklist); Dumas et al. 2010, 7 (distribution); Calor 2011, 320 (checklist).

Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group and most closely resembles A. blahniki and A. hamata in that the apicomesal process of the first article of each inferior appendage is slightly shorter than the second article in these species. It can be recognized by the apical portion of the phallotheca possessing two spines on each “beak”, one apical and the other lateral (Flint 1974). Description. Overall body color brown to yellowish. Frons and vertex of head with brown and whitish setae. Antennal scapes yellowish with elongate dark brown and whitish setae, pedicels yellow, basal flagellomeres light brown and apical flagellomeres brown. First and second articles of maxillary palps with long, brown setae. Legs yellowish to light brown. Forewings brown and irregularly marked with various shades of brown (Flint 1974), covered by small brown and yellow setae, erect dark brown setae of veins, apices with fringes of brown and whitish setae. Sternum V with remarkable anterolateral processes about half as long as sternum (Fig. 2A; Flint 1974). Remarks. Some specimens have 3 spines on each beak of the phallotheca (Fig. 2B), differing from the original description, which cited only 2 spines. The 3rd spine frequently is positioned between the 2 spines described for A. zernyi by Flint (1974) and it has variable length, although it is frequently shorter than the other 2 spines. Parapods also show some variability (Figs. 2C–H). Material examined. BRAZIL: Minas Gerais, Malacacheta, Córrego das Águas Pretas & tribs. Ca. 15 Km S Aiuruoca, 22°03.704’S, 44°38.241’W, el. 1386 m, 21.xi.2001, Holzenthal, R.W., Paprocki, H., Blahnik, R. & Neto; Det. Blahnik, R., 2002, UMSP 000081962 to 000081965, 2 males, 2 females (pinned, MZSP); Rio de Janeiro, Eng. Passos, BR-485, Km 11, Hotel Faz. Palmital 22°25’26”S, 44°44’21”W, el. 960 m, 26.iv.2007, light, Rafael,


GOMES & CALOR

FIGURE 2. Atopsyche (Atopsaura) zernyi Flint, male. 2A, genitalia, left lateral, proctiger and phallotheca omitted, with inset of apex of right inferior appendage, mesal; 2B, phallic apparatus, left lateral. 2C–H—parapod variation, left lateral. fil = filipod.

ATOPSYCHE OF BRAZIL


65

J.A. & Xavier, F., 1 male (pinned, UFBA); Macaé de Cima, Rio Macaé, 22°23684’S, 42°30.136’W, el. 1000 m, 08.iii.2002, Holzenthal, R.W., Blahnik, R., Paprocki, H. & Prather, A.; UMSP 000069095 to 000069120, 1 male, 25 females (pinned, MZSP); same data except Córrego das Flores, 10 Km SE Mury, el. 1000 m, 09.iii.2002, Holzenthal, R.W., Blahnik, R., Paprocki, H. & Prather, A.; Det. Blahnik, R., 2003, UMSP 000088445 to 000088447, 1 male, 2 females (pinned, MZSP); Parati, tributary to Riacho Perequê-açu, 23°12.833’S, 44°47.483’W, el. 190 m, 26.ix.2002, Blahnik, R., Prather, A., Melo, A., Froehlich, C.G. & Silva, R.; Det. Blahnik, R. 2003; UMSP 000087683 to 000087693, 1 male, 10 females (pinned, MZSP); Rio Grande do Sul, São José dos Ausentes, 28°60’407”S, 49°76’236”W, el. 1238 m, 09.iii.2008, Melo, A. & Ferro, V., 11 males (pinned, UFBA); same data except 28°57’319”S, 49°84’292”W, el. 1250 m, 08.iii.2008, 1 male (pinned, UFBA); same data except 28°57’607”S, 49°92’000”W, el. 1250 m, 09.ii.2008, 2 males (pinned, UFBA); same data except 03.ii.2008, 1 male (pinned, UFBA); São Paulo, Salesópolis, Casa Grande, Ribeira, Coruja, 16.xi.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 3 males (alcohol, MZSP); same data except light, 26.i.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 9 males, 6 females (alcohol, MZSP); same data except 16.xi.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 2 males (alcohol, MZSP); same data except 12.xi.1974, Froehlich, C.G.; Det. Flint, O.S. Jr., 1977, 2 males, 1 female (alcohol, MZSP); same data except 12.x.1974, light; Det. Flint, O.S. Jr., 1977, 1 male (alcohol, MZSP); same data except Parque Nacional da Serra da Bocaina, Cachoeira Santo Izidro, 22°44.830’S, 44°36.882’W, el. 1480 m, 02.iii.2002, Blahnik, R. & Paprocki, H.; Det. Blahnik, R. 2003, UMSP 000085844 to 000085852, 1 male, 8 females (pinned, MZSP); Iporanga, Parque Estadual Intervales, 15.xii.1999, light, Bispo, P. & Crisci-Bispo, V., 1 male (alcohol, MZSP); São Luís do Paraitinga, P.E. da Serra do Mar, Núcleo Santa Virgínia, Rio Ipiranga, 15.iv.2009, Mariano, R., Lecci, L. & Schulz, G., 1 male (pinned, UFBA). Distribution. Brazil (Espírito Santo, Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo).

Atopsyche (Atopsaura) diamantina Gomes & Calor, new species Figures 3A–G Diagnosis. This species belongs to the Atopsyche (Atopsaura) longipennis Group, similar to A. iana Mosely 1949, A. atahuallpa Schmid 1989 and A. ayacucho Schmid 1989 in the division of each beak of the phallotheca into two projections, but it differs from these species by the very broad and short parapods, the different shape of the projections on the phallothecal “beaks”, and the broader first and second articles of the inferior appendages. The new species can also be recognized by the upcurved process on the apex of each parapod and by the bifid apex of each “beak” of the phallotheca. Description. Length of each forewing 6.08–6.76 mm (n = 15) long (Fig. 3A). Overall body color yellowish brown, slightly darker dorsally. Frons and vertex of head with erect yellow setae. Antennal scapes brown with long brown setae; pedicels yellowish brown, basal flagellomeres yellow, apical flagellomeres brown. Maxillary palp segments 1 and 2 with long, brown, stout setae apically, labial palps with long, brown setae. Dorsum of thorax with long, erect brown and yellow setae. Legs yellowish brown; coxae of all legs and foreleg tibiae darker brown. Forewings light brown, each with erect brown setae between Cu2 and Cu1A; dark brown setae along costal margin and dark brown patch in middle of wing; apex of wing with fringe of dark brown setae. Tergum III anterolaterally with pair of rounded glands. Sterna II–VII with minute setae. Sterna VI–VII each with prominent spine-like ventral process on posteromesal margin; process on sternum VI long, slender, curved, with short spine-like setae along ventral margin and terminating with longer, spine-like seta; process on sternum VII short, straight. Male genitalia. Segment IX, in lateral view, relatively short longitudinally. Proctiger, in lateral view, broadly widened apically, covered externally with numerous setae. Parapods moderately inflected at middle in dorsal view, apicodorsally each with upcurved hook (Figs. 3C–D). Filipods absent. Preanal appendages relatively large for this genus, setose, capitate, each with short and slender “neck” attached to segment IX and “head” almost square in lateral view (Fig. 3C). Inferior appendages, in lateral view, each with first article broad, slightly narrower basally and apically, its apicomesal lobe thick, digitate (Fig. 3C); second article, in lateral view, thick, wider basally and more slender apically, slightly broader than apicomesal lobe of first article (Fig. 3C); in ventral view, broader than first and acute at apex (Fig. 3G). Phallic apparatus relatively simple; phallotheca broadly rounded basally with dorsal and ventral posterior projections; as viewed laterally, each “beak” of phallotheca divided apically into upcurved dorsal projection and slender, tiny ventral projection (Fig. 3E); in dorsal view, apex of phallotheca


GOMES & CALOR

FIGURE 3. Atopsyche (Atopsaura) diamantina Gomes & Calor, new species, male. 3A, right forewing, dorsal; 3B, right hind wing, dorsal; 3C, genitalia, left lateral, phallic apparatus omitted; 3D, left parapod and preanal appendage, dorsal; 3E, phallic apparatus, left lateral; 3F, phallic apparatus, dorsal; 3G, left inferior appendage, ventral.

ATOPSYCHE OF BRAZIL


67

divided mesally into pair of “beaks”, each beak concave on mesal surface and with posteroventral margin bearing finger-like sclerotized projection (Fig. 3F); aedeagus basally enlarged then spine-like and curving ventrad and caudad. Female and immatures. unknown. Holotype male. BRAZIL: Bahia, Mucugê, Chapada Diamantina, Parque Municipal Sempre Viva, Rio Piabinha, 12°59’34”S, 41°20’27”W, el. 921 m, 25.vii.2010, UV Light Pan Trap, Calor, A.R., Lecci, L., Quinteiro, F., França, D., Arantes, T. & Camelier, P. (alcohol; MZSP). Paratypes. BRAZIL: Bahia, same data as holotype except 2 males (alcohol, UFBA); same data except Rio Cumbuca, 27.vi.2010, UV Light Pan Trap, LEAq team., 1 male (alcohol, UFBA); same data except 12°59’57”S, 41°20’50”W, el. 909 m, 25.vii.2010, light, 1 male (pinned, UFBA); same data except Rio Cumbuca, 27.vi.2010, UV Light Pan Trap, Calor, A.R., Lecci, L., Quinteiro, F., França, D., Arantes, T. & Camelier, P., 1 male (alcohol, UFBA); same data except Andaraí–Igatú, Rio Coisa Boa, 12°53’33.7”S 41°18’58.3”W, el. 664 m, 11.iii.2011, light, Calor, A.R., Camelier, P. & Zanata, A., 1 male (alcohol, UMSP); same data except Rio Coisa Boa, 12°53’13.9”S, 41°18’58.3”W, 12.v.2010, UV Light Pan Trap, França, D., 1 male (alcohol; UFBA); same data except Palmeiras, Vale do Capão, Rio Carinho (ponte), 29.vi.2011, Calor, A.R., Camelier, P. & Burguer, R., 3 males (alcohol, UFBA); same data except Palmeiras, Vale do Capão, Riachinho (ponte), 12°34’19.2”S 41°30’52.5”W, el. 918 m, 25.vi.2011, UV Light Pan Trap, Calor, A.R., Camelier, P. & Burguer, R., 15 males (alcohol, UFBA); same data except 8 males (alcohol, UMSP); same data except Lençóis, Rio Mucugezinho, 12°27’44”S, 41°25’1”W, el. 706 m, 01.viii.2010, UV Light Pan Trap, Calor, A.R., Lecci, L., Quinteiro, F., França, D., Arantes, T. & Camelier, P., 4 males (alcohol, UFBA); same data except Piatã, Fazenda Machado, riacho Machado, cachoeira Caxiban, 13°09’33”S, 41°47’29”W, 30.vii.2010, Calor, A.R., Lecci, L., Quinteiro, F., França, D., Arantes, T. & Camelier, P., 1 male (pinned, UFBA); same data except Riacho Três Morros, cachoeira Patricio, 13°05’12”S, 41°51’12”W, el. 909 m, 29.vii.2010, entomological net, 2 males (pinned, UFBA); same data except light, 1 male (pinned, UFBA); Minas Gerais, Delfinópolis, 03.i.2007, Amorim, D. & Ribeiro, G., 1 male (alcohol, UFBA). Distribution. Brazil (Bahia, Minas Gerais). Etymology. The specific name, a noun in apposition, refers to the Chapada Diamantina, a mountain range in the central region of the State of Bahia, Brazil, which is the type locality of the new species. The word “Diamantina” in Portuguse means “diamantiferous,” and it is an allusion to the large diamond reservoirs in the region.

Atopsyche (Atopsaura) kamakan Gomes & Calor, new species Figures 4A–G Diagnosis. This new species belongs to the Atopsyche (Atopsaura) longipennis Group according to the shape of the apicomesal process of the first article of each inferior appendage which doubles the second article. The new species resembles to A. huarcu and A. sanctipauli in that the second article of each inferior appendage is broad basally and its apex is slender and curved so as to cover the apicomesal process of the first article (Flint 1974) and in that the phallotheca has an elongate, curved, spine-like basodorsal process. This new species can be recognized by the shape of the anterodorsal lobe of the apicomesal process of the first article of each inferior appendage and by the upturned and uniformly slender basodorsal process of the phallotheca. Description. Length of each forewing 5.52–6.56 mm (n = 19) (Fig. 4A). Overall body color yellow to dark brown. Head, frons and vertex with brown setae. Antennal scapes light yellow with long setae, pedicels light yellow; basal flagellomeres yellow, apical ones brown. Setose labial and maxillary palps with long brown setae. Dorsum of thorax with erect light and dark brown setae. Legs yellowish brown, coxae of all legs and foreleg tibiae dark brown, mid- and hind leg tibiae with yellow and brown. Forewings brown, erect setae of veins forming distinct pattern of alternating dark brown and yellow setae, with fewer, scattered white setae (Fig. 4A). Terga II, III, IV, V, VI and VII with elongate, coarse setae on posterolateral margins; terga III and IV each with pair of prominent, rounded glands located at anterolateral margins of respective terga. Sterna II, III, IV, V and VI with numerous minute setae; sternum VI process long, slender, curved, with longer, spine-like setae; process on sternum VII short, straight.


GOMES & CALOR

FIGURE 4. Atopsyche (Atopsaura) kamakan Gomes & Calor, new species, male. 4A, right forewing, dorsal; 4B, right hind wing, dorsal; 4C, genitalia, left lateral, phallic apparatus omitted, detail of the apex of the inferior appendage in median view; 4D, left parapod and pre-anal appendage, dorsal; 4E, phallic apparatus, left lateral; 4F, phallic apparatus, dorsal; 4G, left inferior appendage, ventral.

ATOPSYCHE OF BRAZIL


69

Male genitalia. Segment IX, in lateral view, longer ventrally tapering dorsally (Fig. 4C). Proctiger as viewed laterally, broadly widened apically, covered externally with numerous minute setae, lateral margins with several elongate setae. Parapods broad at base, narrow at tip, each with apicodorsal process upturned and with posterolateral process subapical; in dorsal view, parapods irregular with complex apex (Fig. 4D). Filipods absent. Preanal appendages small, rounded and setose. Inferior appendages each with first article relatively slender and four times as long as greatest width near mid length, apicomesal projection long and with mesolateral tooth; second article bilobed apically, anterodorsal lobe slender and acute and other lobe setose; in ventral view, second article broad basally and apicolateral margin parallel and closely aligned with the anteromesal projection of first article (Fig. 4G). Phallic apparatus relatively simple, apex divided mesally into pair of lateral “beaks”, in lateral view, “beaks” uniformly wide and rounded apically (Fig. 4E), slender in dorsal view (Fig. 4F); phallotheca with elongate, the upturned and uniformly slender basodorsal process, with dorsal tooth near middle, uniformly slender and apically upturned; aedeagus forming elongate curved spine, with basal sclerite (Fig. 4E). Female and immatures. unknown. Holotype male. BRAZIL: Bahia, Camacan, Reserva Particular do Patrimônio Natural Serra Bonita, 15°23’02”S, 39°34’10”W, iv.2012, Malaise trap, Calor, A.R. et al. (alcohol, MZSP). Paratypes. BRAZIL: Bahia, same as holotype except 04.xi.2009, 1ª cachoeira, UV Light Pan Trap, 2 males (alcohol, UFBA); same as holotype except 05.xi.2009, Centro de Pesquisa, UV Light Pan trap, 2 males (alcohol, UFBA); same as holotype except 26–27.xi.2011, caixa d’água, UV Light Pan Trap, 2 males (alcohol, UFBA); same as holotype except 28.xi.2011, Quinteiro, F., Dias, E. & Duarte, T., 1 male (alcohol, UFBA); same as holotype except Fechadinho, 15°23’9.1”S, 39°34’3.1”W, el. 723 m, 27.xi.2011, UV Light Pan Trap, 1 male (alcohol, UFBA); same as holotype except 15°23’03”S, 39°34’00.1”W, el. 723 m, iii.2009, Malaise trap, 1 male (alcohol, UFBA); same as holotype except 15°23’9.1”S, 39°34’3.1”W, el. 723 m, 29.x.2008, UV Light Pan Trap, Calor, A.R., Mariano, R. & Mateus, S., 1 male (pinned, UFBA); 15°28’01”S, 39°33’56”W, el. 820 m, 01.vii.2008, UV Light Pan Trap, Calor, A.R., Lecci, L., Pinho, L. & Moretto, R., 1 male (alcohol, UFBA); same as holotype except 15°23’28”S, 39°33’56”W, el. 820 m, 28.x.2008, UV Light Pan Trap, Calor, A.R., Mariano, R. & Mateus, S., 1 male (pinned, UFBA); Santa Teresinha, Serra da Jibóia, Pedra Branca, riacho das torres, cachoeira, 12°51’00”S, 39°28’48”W, 28.ix.2009, el. 678m, UV Light Pan Trap, Calor, A.R. & Cruz, A., 1 male (alcohol, UFBA); same data except 12°51’025”S, 39°28’805”W, el. 687 m, 27.x.2008, UV and white light, Calor, A.R., Mariano, R. & Mateus, S., 1 male (pinned, UFBA); same data except UV Light Pan Trap, Calor, A.R., Quinteiro, F.; França, D., Mariano, R. & Costa, A., 1 male (alcohol, UFBA); same data except waterfall, 12°51’00”S, 39°28’48”W, 04.ii.2010, 1 male (alcohol, UFBA); same data except 06.x.2010, UV Light Pan Trap, Calor, A.R., Quinteiro, F., França, D., Mariano, R. & Costa, A.M., 1 male (alcohol, UFBA); same data except 08.xi.2010, 1 male (alcohol, UFBA); Elísio Medrado, Serra da Jibóia, Jequitibá reserve, 12°52’215”S, 39°28’565”W, 07.xi.2010, Silva-Neto, A. & Araújo, T., 1 male (alcohol, UFBA); same data except Wenceslau Guimarães, Estação Ecológica Wenceslau Guimarães, sede, riacho Serra Grande, 13°35’34”S, 39°42’52”W, 10.x.2010, UV Light Pan Trap, Calor, A.R., Quinteiro, F., Costa, A., 2 males (alcohol, UFBA); same data except 09.x.2010, 1 male (alcohol, UMSP). Distribution. Brazil (Bahia). Etymology. The Kamakã language (Kamakan), or Ezeshio, is an extinct language of a small family believed to be part of the Macro-Gê languages of Bahia State, near Brazil's Atlantic coast. Dialects included Kotoxó, Mongoyó/Mangaló, and Kamakã-Menien. The Kamakan native indigenous people lived in the neighbourhoods of the distribution area of the new species, now a municipality known as Camacan.

Atopsyche (Atopsaura) muelleri Gomes & Calor, new species Figures 5A–G Diagnosis. This new species belongs to Atopsyche (Atopsaura) longipennis Group according to the shape of the plate of the apicomesal margin of the first article of each inferior appendage which doubles the second article. Atopsyche muelleri new species is similar to A. serica and A. huanapu in the broad segment IX, the general shape of the parapods, in short filipods, and in the concave mesal face of the second article of each inferior appendage. Moreover, it resembles A. siolii in the two apical projections of the phallotheca, but the species can be recognized by the concave mesal face of the second article of each inferior article and by the bilobed apex of the phallotheca.


GOMES & CALOR

FIGURE 5. Atopsyche (Atopsaura) muelleri Gomes & Calor, new species, male. 5A, right forewing, dorsal; 5B, right hind wing, dorsal; 5C, genitalia, left lateral, phallic apparatus omitted; 5D, left parapod and pre-anal appendage, dorsal; 5E, phallic apparatus, left lateral; 5F, phallic apparatus, dorsal; 5G, left inferior appendage, ventral.

Description. Length of each forewing 6,09–6.75 mm (n = 6) (Fig. 5A). Overall body color yellow to light brown. Frons and vertex of head with stout, erect brown and light yellow setae. Antennal scapes brown with brown setae, pedicels light brown; basal flagellomeres yellow, apical flagellomeres brown. Maxillary palps absent ATOPSYCHE OF BRAZIL


71

(broken), setae of labial palps yellow and brown. Thorax with light yellowish and stout brown setae. Legs yellowish brown, coxae of all legs and foreleg tibiae darker brown, mid- and hind leg tibiae with light yellow setae. Forewings brown, erect setae of veins forming irregular pattern of alternating dark brown and yellow setae (Fig. 5A). Tergum II bearing pair of prominent glands at posterolateral margins; tergum III with pair of prominent glands at posterolateral margins. Sternum V with pair of small, convexly rounded glands, sterna VI and VII each with prominent spine-like ventral process on posteromesal margin; process on VI long, slender, curved with short spinelike setae along ventral margin and terminating with longer, spine-like setae. Male genitalia. Segment IX long laterally. Proctiger, in lateral view, broadly widened apically, with angulated apical margin, covered with minute setae. Parapods narrow, relatively simple, in lateral view rounded apicoventrally, each with acute, setose and upturned apicodorsal projection (Fig. 5C); in dorsal view, parapods with relatively uniform width (Fig. 5D). Preanal appendages small, rounded and setose. Filipods short, slender, without setae, slightly longer than preanal appendages. Inferior appendages each with first article long, three times as long as wide, slightly narrower at base, with subtriangular apicomesal projection; second article broad mesally and narrowed at tip (Fig. 5C), concave on mesal face (Fig. 5G). Phallic apparatus relatively simple; phallotheca in lateral view with beaks elongate and forked apically in lateral view; ventral lobe short and rounded (Fig. 5E). Phallotheca in dorsal view with beaks slightly sinuous and broadened subapically, apically pointed with apices directed mesad (Fig. 5F). Female and immatures. unknown. Holotype male. BRAZIL: Goiás, Chapada dos Veadeiros, Ribeirão Água Fria, 16.xii.2006, Bispo, P., Yokoyama, E. & Paciencia, G. (alcohol, MZSP). Paratypes. BRAZIL: Goiás, same as holotype except Ribeirão Raizama, 10.xii.2006, light, Bispo, P., Yokoyama, E. & Paciencia, G., 2 males (alcohol, MZSP), 3 males (alcohol, UFBA). Distribution. Brazil (Goiás). Etymology. This new species is named in honor of Fritz Müller, the “prince of observers,” according to Charles Darwin. The naturalist Fritz Müller was a pioneer in studies of Neotropical caddisflies (among other taxa) and an important collaborator and contributor to the argumentation for Darwin’s Evolution Theory, especially by means of Müller’s book “Für Darwin” (1864) and its translations.

Key to males of Atopsyche species from Brazil 1 2(1) 3(2)

First article of each inferior appendage without apicomesal projection (subgenus Atopsyche) . . . . . . . . . . . . . . . . . . . . . . . . 2 First article of each inferior appendage with apicomesal projection (subgenus Atopsaura (A.) longipennis Group) . . . . . . . . 5 Second article of each inferior appendage tapered from base and hooked ventrad (A. bolivari Group) . . . . . . . . . . . . . . . . . . . 3 Second article of each inferior appendage broadened at base and trianguloid (A. tripunctata Group) . . . . . . . . . . . . . . . . . . . 4 Posterolateral beaks of phallotheca posteriorly each divided into two long, paired processes, dorsal process longer and bifurcate apically, ventral process curved mesad and crossing process from opposing beak (Santos & Holzenthal 2012, fig. 2D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. parauna Posterolateral beaks of phallotheca posteriorly wide, truncate or shallowly divided, with apicoventral corners downturned (Santos & Holzenthal 2012, fig. 2E–G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. urumarca 4(2) Second article of each inferior appendage evenly tapering and rounded at apex (Ross & King 1952, fig. 3A) . . . . . . A. erigia Second article of each inferior appendage obtuse and truncate at apex (Schmid 1989, pl. XVI, fig. 1) . . . . . . . . A. chirihuana 5(1) Filipods absent (Fig. 1A). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Filipods present (Fig. 2A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 6(5) Phallotheca without dorsal process (Fig. 3E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Phallotheca with dorsal process (Fig. 1B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 7(6) Apical portion of first article of each inferior appendage bifurcate (Fig. 4C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Apical portion of first article of each inferior appendage not bifurcate (Schmid 1989, pl. XVII, fig. 5) . . . . . . . . . . . . . . . . . 10 8(7) Basodorsal process of phallotheca elongate, curved, spine-like (Fig. 4E) . . . . . . . . . . . . . . . . . . . . . . . A. kamakan new species Basodorsal process of phallotheca broadening at tip (Flint 1974, fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9(8) Beaks of phallotheca each with lateral thin, deeply bifurcate lobe at mid-length (Flint 1974, fig. 27) . . . . . . . . . A. sanctipauli Beaks of phallotheca without lateral thin, deeply bifurcate lobes (Schmid 1989, pl. XVII, fig. 9) . . . . . . . . . . . . . . . . A. huarcu 10(7) Basodorsal process of phallotheca longer than inferior appendages (Schmid 1989, pl. XVII, fig. 5) . . . . . . . . . A. huamachucu Basodorsal process of phallotheca shorter than inferior appendages (Schmid 1989, pl. XVII, fig. 3). . . . . . . . . . . . . . . . . . . 11 11(10) Basodorsal process of phallotheca tapering at tip (Schmid 1989, pl. XVII, fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . A. huacachaca Dorsal process of phallotheca broadening at tip (Schmid 1989, pl. XVII, fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . A. hatunpuna 12(6) Inferior appendages each with apicoventral process (Marlier 1964, fig. 3A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. plancki


GOMES & CALOR

13(12) 14(5) 15(14) 16(15) 17(16) 18(15) 19(14) 20(19) 21(19) 22(21) 23(21) -

Inferior appendages without apicoventral processes (Figs. 3C, 3G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Parapods forked and terminating in two slender branches (Schmid 1989, pl. XVIII, fig. 10) . . . . . . . . . . . . . . . . . A. apurimac Parapods not forked, but apicodorsally with upcurved sclerotized projection (Fig. 3C) . . . . . . . . . . A. diamantina new species Filipods shorter than parapods (Fig. 2A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Filipods longer or about as long as parapods (Santos & Holzenthal 2012, fig. 1A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Inferior appendages downcurved at midlength (Ross 1953, fig. 9A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Inferior appendages straight (Schmid 1989, pl. XVII, fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Short thumb-like process present mesoventral of filipods and parapods and above phallotheca (Ross 1953, fig. 9A) … A. serica Thumblike process absent (Fig. 5C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Phallic apparatus with middorsal hood (Flint 1971, fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. siolii Phallic apparatus without middorsal hood (Fig. 5D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. muelleri new species Second article of each inferior appendage pyriform; parapods about as long as basal height, abruptly taller in apical third (Schmid 1989, pl. XVIII, fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. acahuana Second article of each inferior appendage only slightly larger basally, not pyriform; parapods four times longer than tall, each with dorsal margin forming slight discontinuity in apical third (Schmid 1989, pl. XVIII, fig. 6) . . . . . . . . . . . . . . . A. antisuya Filipods about as long as parapods (Ross & King 1952, fig. 21A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Filipods longer than parapods (Santos & Holzenthal 2012, fig. 1A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Beaks of phallotheca each bearing short, strongly sclerotized posteroventral spine beneath long, setose, upturned process (Ross & King 1952, fig. 21B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. hamata Thumb-like process absent mesoventral of filipods and parapods (Ross & King 1952, fig. 22A). . . . . . . . . . . . . A. longipennis Phallotheca with long paired processes (Santos & Holzenthal 2012, fig. 1E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Phallotheca without long paired processes (Schmid 1989, pl. XVII, fig.14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Beaks of phallotheca with several spines on their lateral edges; filipods exceeding length of parapods (Santos & Holzenthal 2012, fig. 1A, 1B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. blahniki Beaks of phallotheca without spines on their lateral edges (Fig. 2B; Flint 1974, fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . A. zernyi Second article of each inferior appendage dactylate (Denning & Sykora 1968, fig. A) . . . . . . . . . . . . . . . . . . . . . . . A. usingeri Second article of each inferior appendage concave on mesal face (Schmid 1989, pl. XVII fig. 14) . . . . . . . . . . . . A. huanapu

Acknowledgements We especially thank Dr. Paola Rueda Martín for giving valuable comments and corrections on this manuscript. The authors thank Dr. Roger Blahnik for helpful comments on diagnostic characters. We thank the Laboratório de Entomologia Aquática (LEAq, UFBA) for field assistance. We are also grateful to Dr. Maria Teresa Stradmann, in memorian (Grupo Ambientalista da Bahia, GAMBÁ), Dr. Vitor Becker and Sra. Clemira Souza (Reseve Serra Bonita, Instituto Uiraçú) for assistance in field. The authors also thank the curators Dr. Freddy Bravo (Universidade Estadual de Feira de Santana), Dr. Sonia Casari, Dr. Carlos Campaner (Museu de Zoologia da Universidade de São Paulo) for the loan of the biological material, Dr. Elisa Angrisano and Dr. Paola R. Martín for support during the analysis of specimens in Argentina. We also thank Chico Mendes Institute for Biodiversity Conservation (ICMBio) for issuing collecting permits. This work was supported by Foundation for Research Support of the State of Bahia (FAPESB) grant (process 5716/2009), by National Council for Scientific and Technological Development (CNPq) grant (processes 558317/2009-0, 473703/2010-6, 552525/2010-3). VG and ARC thank CNPq fellowships (135585/2011-2, 141367/2004-0), SWE 201382/2007-5, 243238/2014). We highly appreciate the valuable suggestions and corrections of the editor, Dr. John Morse, and reviewers on our manuscript.

References Banks, N. (1905) Description of new Neartic neuropteroid insects. Transactions of the American Entomological Society, 32, 1– 20. Banks, N. (1913) Synopses and descriptions of exotic Neuroptera. Transactions of the American Entomological Society, 39, 201–242. Blahnik, R.J. & Gottschalk, R.M. (1997) New species of Atopsyche from Costa Rica (Trichoptera: Hydrobiosidae). Entomological News, 108 (3), 161–174. Blahnik, R.J., Paprocki, H. & Holzenthal, R.W. (2004) New distribution and species records of Trichoptera from southern and southeastern Brazil. Biota Neotropica, 4 (1), 1–6. http://dx.doi.org/10.1590/S1676-06032004000100009 Bueno-Soria, J. & Flint, O.S. Jr. (1980) [for 1978]. Catálogo sistemático de los tricópteros de México (Insecta: Trichoptera), con algunos registros de Norte, Centro y Sudamérica. Anales del Instituto de Biologia Universidad Nacional Autonoma de

ATOPSYCHE OF BRAZIL


73

Mexico, Serie Zoologia, 49 (1), 189–218. Calor, A.R. (2011) Checklist dos Trichoptera (Insecta) do Estado de São Paulo, Brasil. Biota Neotropica, 11, 318–328. http://dx.doi.org/10.1590/S1676-06032011000500028 Calor, A.R. & Mariano, R. (2012) UV light pan traps for collecting aquatic insects. Entomobrasilis, 5 (2), 164–166. http://dx.doi.org/10.12741/ebrasilis.v5i2.187 Curtis, J. (1835) British Entomology. XII. Richard Taylor, London, 530–577. Denning, D.J & Sykora, J.L. (1968) Three new species of Trichoptera from Brazil. Beiträge zur Neotropischen Fauna, 5, 172– 177. http://dx.doi.org/10.1080/01650526809360405 Dumas, L.L., Santos, A.P.M., Jardim, G.A., Ferreira, N.F. Jr. & Nessimian, J.L. (2010) Insecta, Trichoptera: New records from Brazil and other distributional notes. CheckList, 6 (1), 7–9. Edwards, S.W. & Arnold, C.R. (1961) The caddisflies of the San Marcos River. The Texas Journal of Science, 13, 398–415. Flint, O.S. Jr. (1971) Studies of neotropical caddisflies, XII: Rhyacophilidae, Glossosomatidae, Philopotamidae, and Psychomyiidae from the Amazon Basin (Trichoptera). Amazoniana, 3 (1), 1–67. Flint, O.S. Jr. (1972) Studies of neotropical caddisflies, XIV: On a collection from northern Argentina. Proceedings of the Biological Society of Washington, 85 (17), 223–248. Flint, O.S. Jr. (1974) Studies of neotropical caddisflies, XVIII: New species of Rhyacophilidae and Glossosomatidae (Trichoptera). Smithsonian Contributions to Zoology, 169, i–iv, 1–30. http://dx.doi.org/10.5479/si.00810282.169 Flint, O.S. Jr., Holzenthal, R.W. & Harris, S.C. (1999) Catalog of Neotropical Caddisflies (Insecta: Trichoptera). Ohio Biological Survey, Columbus, Ohio, iv + 239 pp. Holzenthal, R.W., Blahnik, R.J., Prather, A.L. & Kjer, K.M. (2007) Order Trichoptera Kirby, 1813 (Insecta), caddisflies. Zootaxa, 1668, 639–698. Holzenthal, R.W. & Cressa, C. (2002) The Trichoptera, caddisflies, of Venezuela: Three new species and records of Atopsyche Banks (Hydrobiosidae). Studies on Neotropical Fauna and Environment, 37 (2), 133–143. http://dx.doi.org/10.1076/snfe.37.2.133.8578 Holzenthal, R.W., Morse, J.C. & Kjer, K.M. (2011) Order Trichoptera Kirby, 1813. In: Zhang, Z-Q. (Ed.), Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa, 3148, 209–211. Marlier, G. (1964) Sur trois Trichopteres nouveaux recueillis en Amerique du Sud par le Professeur J. Illies. Bulletin Institut Royal des Sciences naturelles de Belgique, 40 (6), 1–15. McLachlan, R. (1866) Descriptions of new or little-known genera and species of exotic Trichoptera; with observations on certain species described by Mr. F. Walker. Transactions of the Entomological Society of London, Series 3, 5, 247–278. McLachlan, R. (1868) On some new forms of trichopterous insects from New Zealand; with a list of the species known to inhabit those colonies. The Journal of the Linnean Society, Zoology, 10, 196–214. Mosely, M.E. (1949) New Trichoptera and a redescription of Leptocellodes flaveola Ulmer. Proceedings of the Royal Entomological Society of London, 18 (B), 37–41. Müller, F. (1864) Für Darwin. Verlag von Wilhelm Engelmann, Leipzig [(1869) translated into English as “Facts and Arguments for Darwin,” 152 pp.; (2009) translated into Portuguese as “Para Darwin,” Editora da Universidade Federal de Santa Catarina, Santa Catarina, 280 pp. Müller, F. (1921) Briefe und noch nicht veröffentliche Abhandlungen aus dem Nachlass 1854-1897. In: A. Möller (Ed.), Fritz Müller: Werke, Briefe und Leben, 2. G Fischer, Jena, pp. 383–642. Navás, L. (1924) Insectos de la América Central. Brotéria, Série Zoológica, 21, 55–86. Paprocki, H., Blahnik, R.J. & Holzenthal, R.W. (2004) Checklist of the Trichoptera (Insecta) of Brazil. Biota Neotropica, 4 (1), 1–22. http://dx.doi.org/10.1590/S1676-06032004000100008 Parkinson, J. (1811) Organic Remains of a Former World. An Examination of the Mineralized Remains of the Vegetables and Animals of the Antidiluvian World; Generally Termed Extraneous Fossils. The Third Volume; Containing the Fossil Starfishes, Echini, Shells, Insects, Amphibia, Mammalia. Sherwood, London, xv + 479 pp. Reynaga, M.C. & Rueda Martín, P. (2010) Trophic analysis of two species of Atopsyche (Trichoptera: Hydrobiosidae). Limnologia, 40, 61–66. http://dx.doi.org/10.1016/j.limno.2008.07.004 Ross, H.H. (1947) Descriptions and records of North American Trichoptera, with synoptic notes. Transactions of the American Entomological Society, 73, 125–168. Ross, H.H. (1951a) The origin and dispersal of a group of primitive caddisflies. Evolution, 5 (2), 102–115. http://dx.doi.org/10.2307/2405762 Ross, H.H. (1951b) Relationships of the Fijian species of Apsilochorema (Trichoptera, Rhyacophilidae). Occasional Papers of Bernice P. Bishop Museum, Honolulu, Hawaii, 20 (12), 175–182. Ross, H.H. (1953) Additional material on the phylogeny and dispersal of Atopsyche (Trichoptera, Rhyacophilidae). Journal of the Washington Academy of Sciences, 43, 287–293. Ross, H.H. (1956) Evolution and Classification of the Mountain Caddisflies. University of Illinois Press, Urbana, 213 pp. Ross, H.H. & King, E.W. (1951) The caddisfly tribe “Hydrobiosini” (Trichoptera, Rhyacophilidae). Acta Zoologica Lilloana,


GOMES & CALOR

12, 501–508. Ross, H.H. & King, E.W. (1952) Biogeographic and taxonomic studies in Atopsyche (Trichoptera, Rhyacophilidae). Annals of the Entomological Society of America, 45 (2), 177–204. http://dx.doi.org/10.1093/aesa/45.2.177 Santos, A. & Holzenthal, R.W. (2012) Three new species of Atopsyche Banks (Trichoptera, Hydrobiosidae) from Brazil. ZooKeys, 207, 35–78. http://dx.doi.org/10.3897/zookeys.207.3419 Schmid, F. (1970) Le genre Rhyacophila et la famille de Rhyacophilides. Mémoirs de la Société du Canada, 66, 1–230. Schmid, F. (1989) Les Hydrobiosides (Trichoptera, Annulipalpia). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, 59 (Supplement), 1–154. Stephens, J.F. (1835–1837) Illustrations of British Entomology or a Synopsis of Indigenous Insects: Containing their Generic and Specific Distinctions; with an Account of their Metamorphoses, Times of Appearance, Localities, Food, and Economy, as far as Practicable. Mandibulata, Vol. VI. Baldwin and Cradock, London, 240 pp. [Trichoptera, pp. 146–208, 1836; 209–234, 1837]. Ulmer, G. (1905) Über die geographische Verbreitung der Trichopteren. Zeitschrift für wissenschaftliche Insektenbiologie, 1, 16–32, 68–80, 119–126. Ulmer, G. (1907) Trichoptera. In: Wytsman, P.A.G. (Ed.), Genera Insectorum, part 60a, L. Desmet Verteneuil, Bruxelles, pp. 1– 259, pls. 1–41. Ulmer, G. (1909) Argentinische Trichopteren. Zeitschrift für wissenschaftliche Insektenbiologie, 5, 73 –76, 120–124. Ulmer, G. (1957) Köcherfliegen (Trichopteren) von den Sunda Inseln. Teil III. Archiv für Hydrobiologie, 23 (Supplement), 109–470. Walker, F. (1852) Catalogue of the Specimens of Neuropterous Insects in the Collection of the British Museum, Part I: Phryganides-Perlides. British Museum, London, 192 pp. Ward, J.B., Leschen, R.A.B., Smith, B.J. & Dean, J.C. (2004) Phylogeny of the caddisfly (Trichoptera) family Hydrobiosidae using larval and adult morphology, with the description of a new genus and species from Fiordland, New Zealand. Records of the Canterbury Museum, 18, 23–43. Wiggins (2004) Caddisflies: The Underwater Architects. University of Toronto Press, Toronto, 292 pp.

ATOPSYCHE OF BRAZIL


75