Trichoptera: Hydropsychidae)

ANNALES HISTORICO-NATURALES MUSEI NATIONALIS HUNGARICI Volume 104 Budapest, 2012 pp. 215–297

New species and records of Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)

J. OLÁH1 & K. A. JOHANSON2

1H-4032 Debrecen, Tarján u. 28, Hungary. Email: [email protected] 2Swedish Museum of Natural History, Entomology Department, Box 50007, S-104 05 Stockholm, Sweden. Email: [email protected]

– Fifty-four new species in the hydropsychid genera Centromacronema ULMER, 1905 (3 new species), Leptonema GUÉRIN-MÉNEVILLE, 1843 (10 new species), Macronema PICTET, 1836 (2 new species), Macrostemum KOLENATI, 1859 (1 new species), and Smicridea MCLACHLAN, 1871 (38 new species) are described and illustrated. New records of 48 species in the above listed genera, as well as Plectromacronema ULMER, 1906, are given. With 287 figures and 1 table.

– Hydropsychidae, new species, Neotropical Region, new records.

INTRODUCTION

According to FLINT et al. (1999) about 370 species of Hydropsychidae CURTIS, 1835 are known from the Neotropical Region. Only slightly more species were listed by MORSE (2012). The family is one of the most diverse in the neotropics, surpassed only by the Hydroptilidae STEPHENS, 1836. The Hydropsychidae are divided into the subfamilies Diplectroninae ULMER, 1951, Hydropsychinae CURTIS, 1834, Macronematinae ULMER, 1905 and Smicrideinae FLINT, 1974. The Arctopsychidae are considered a subfamily of Hydropsychidae by some authors (HOLZENTHAL et al. 2007, MORSE 2012), and regarded as a distinct family by others (e.g. SCHMID 1968). In the Neotropical Region, the Macronematinae and Smicrideinae comprise 90% of the known species diversity in the family. These two groups cover the study organisms of this work.

Annls hist.-nat. Mus. natn. hung. 104, 2012 216 J. Oláh & K. A. Johanson

MATERIAL AND METHODS

The study is based on 1495 males and 1749 females belonging to 102 species in 6 genera. All material is preserved in 70–80% alcohol. In order to observe morphological de- tails in the genitalia, the entire abdomen was removed and placed in a small glass beaker of 25 cm3 with 10% KOH solution and boiled for 5–15 minutes for digestion. The duration of the treatment is adjusted individually to the effectiveness of clearing process, which de- pends on the species or even on the nutritive state of tissues or on the physiological condi- tion of the specimens. The process of digestion was followed by examining the transparency of the specimen under treatment, as the dissolution rate of the soft tissues is clearly visible to naked eye. The digested abdomen was subsequently transferred to distilled water and the macerated tissue was removed mechanically by fine tipped forceps and needles. The cleared abdomen was transferred to 80% ethyl alcohol, and to glycerine for examination under microscope. Different sized pins modified to supporting ring bottom was intro- duced into the abdomen and used to hold and stabilise the genitalia in lateral, dorsal and ventral position for drawing. However, the plane of view is never perfect and we made no special procedures of grid, matrix or reflection to produce absolute mirror symmetry of the drawings. Instead, the genital structures are drawn exactly as seen in the microscope. How- ever setae are represented only by their alveoli and moreover their density is only symbolic. If essential the setal length or setal shape are presented by drawing a single or a few setae only. The genital structure was traced by pencil on white paper using a drawing tube moun- ted on a WILD M3Z microscope at between 260–416× magnification. Final illustrations were prepared by enlarging the original pencil drawings and re-drawn on transparent paper by Black India Ink. The inked illustrations were scanned on an Epson Expression 1680 Pro scanner in grayscale and 800 dpi resolution. The plates were arranged, and brightness and contrast edited in Adobe® Photoshop© 8.0 on a Macintosh G5.

We used our functional appendicular terminology and not the conventional anato- mical directional terminology to describe the genital structures in species description (OLÁH &JOHANSON 2008). Species descriptions were standardised to ensure consistently formatted and comparable description in general accord with EVENHUIS’ (2007) template principle. We have standardised also the terminology to describe space extensions of variously formed structural elements. The following terms were used to qualify the dimensions and extensions of genital structural elements: (1) short or long for length dimension on the longitudinal direction of coronal plane along the anteroposterior axis; (2) low or high (traditionally shallow or deep especially for excisions) for height dimension on the vertical direction of the sagittal plane along the dorsoventral axis and (3) narrow or wide (broad)on the lateral direction of the transversal plane along the mediolateral or left-right axis. The three dimensional Cartesian coordinate system provides theoretical possibility to quantify by measurements the three physical dimensions of length, width, and height of each structural element. However, this quantification is used very seldom in species description. Here we quantify only the length of forewing.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)217

The material is deposited in the following collections: HNHM = Hungarian Natural History Museum, Budapest, Hungary; NRM = Naturhistoriska Riksmuseet, Entomology Department, Stockholm, Sweden; OPC = OLÁH Private Collection, Debrecen, presently under protection of the HNHM.

TAXONOMIC PART

CURTIS, 1834 Centromacronema ULMER, 1905 Centromacronema auripenne RAMBUR, 1842

Material examined – : Approuaguekaw, Kaw Mt, 4°32.833’N, 52° 11.452’W, 77 mao, 24.I.2007, FRG 5, light trap, leg. N. JÖNSSON (1 female, NRM).

Remarks – Forewing nearly uniform brownish; crossvein m present, producing closed median cell.

Centromacronema excisum ULMER, 1905

Material examined – : San Martin Prov., creek crossing rd. Juan Guerra-Cha- zuta, 14 km (rd.) E Colombia Bridge, 6°35.594’S, 76°13.172’W, light, loc. 09, 9.I.2009 leg. K. A. JOHANSON & T. MALM (1 female, NRM).

sp. n. (Figs 1–5)

Diagnosis – This species is distinguished from most other Centromacronema by the loss of crossvein m, leaving the median cell open on forewing. Its upward directed dorsoapical corner of the phallic apex distinguish this new species from all other known species.

Description – Male. Dorsum of head, thorax and abdomen dark brown, light brown or even yellow below; forewing brown without distinct pattern; there is some indistinct paler area between Sc and M; forewing length 8.2 mm; crossvein m lost on forewing. Maxillary palp formula 1-2-4-3-5, segment 3 as long as 1, 2 and 4 together. Tibial spurs 044, dorsoapical projection from foretibia pointed accompanied by 4-5 small spines. Cuticular structure of sternum V glands forming a pair of rounded strongly sclerotised ridge. Male genitalia. Segment IX short, its dorsum with excised apex slightly roofing over segment X; excision varying in paratypes from small to none; intersegmental profile between segments IX and X high obtuse angled; stout setae present on apical rim. Segment

Annls hist.-nat. Mus. natn. hung. 104, 2012 218 J. Oláh & K. A. Johanson

X high basally and very low apically with pointed apex in lateral view; cerci located middle as a pair of small elevated setose warts; vestigial paraproct developed as a pair of ventrolateral straps directed downward from apical lobes. Coxopodite and harpago fused, slightly S-forming. Phallic organ with dorsoapically produced apex in lateral view; with very long and very narrow mesal excision in dorsal view; phalloteremal sclerites with specific pattern.

Material examined – Holotype, male: : River Salvation, Mt Manu, Mother of Good, 546 m, humid subtropical forest, 71°21’22’’W, 12°50’16’’S, 26–28.II.2005, Malaise trap, leg. JUAN CHAVEZ LOPEZ (NRM). Paratypes: same data as holotype (4 males, NRM, 3 males, OPC).

Etymology – Felfele, from “felfelé”, upward in Hungarian, refers to the upward produced dorsoapical corner of the phallic apex in lateral view.

1 5 4

Centromacronema felfele sp. n., holotype, male genitalia: 1 = lateral view; 2 = left gonopod, ventral; 3 = dorsal; 4 = phallus lateral; 5 = phallus apex, ventral

sp. n. (Figs 6–10)

Diagnosis – This species is distinguished from most other Centromacronema by the loss of crossvein m, leaving the median cell open. It resembles C. auripenne (RAMBUR, 1842) from Brazil, from which it differs by having median cell open on forewing and differently shaped phallic apex.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)219

Description – Male. Dorsum of head, thorax and abdomen dark brown, forewing brown without distinct pattern; there is some indistinct paler area between Sc and M; forewing length 16 mm; crossvein m lost on forewing. Maxillary palp formula 1-(2,4)-3-5, segment 3 as long as 1, 2 and 4 together. Tibial spurs 044, dorsoapical projection from foretibia pointed. Cuticular structure of sternum V glands forming a pair of auriform strongly sclerotised ridge. Male genitalia. Segment IX short, its dorsum with excised apex slightly roofing over segment X; intersegmental profile between segments IX and X high obtuse angled; stout setae present on apical rim. Segment X high basally and very low apically with pointed apex in lateral view; cerci located middle as a pair of elevated setose warts; vestigial paraproct developed as a pair of ventrolateral straps directed downward from apical lobes. Coxopodite and harpago fused, slightly S-forming, constricted middle and capitate. Phallic organ with spatulate apex in lateral view; with long and narrow mesal excision in dorsal view; phallotremal sclerites with specific pattern.

8 9 7

Centromacronema kanalas sp. n., holotype, male genitalia: 6 = lateral view; 7 = left gonopod, ventral; 8 = dorsal; 9 = phallus lateral; 10 = phallus apex, ventral

Material examined – Holotype, male: : dept. Pasco, Yanachaga Chemilien NP, INRENA Refugio El Cedro, S10°32.717, W75°21.492, 2460 m, 30.I.2003, leg A. KUN &B. BENEDEK, HNHM expedition (HNHM).

Etymology – Kanalas, from “kanalas”, spatulate in Hungarian, refers to the shape of the phallic apex dorsal view.

Annls hist.-nat. Mus. natn. hung. 104, 2012 220 J. Oláh & K. A. Johanson

sp. n. (Figs 11–15)

Diagnosis – It is closest to C. auripenne (RAMBUR, 1842), but differs by having crossvein m lost on forewing and apex of tergite IX not excised.

Description – Male. Dorsum of head, thorax and abdomen dark brown, forewing brown without distinct pattern, except some lighter yellowish basal half and there is some paler area between Sc and M; yellowish basal half indistinct or variously distinct in paratypes from the same habitat; forewing length 9.5 mm; crossvein m lost on forewing. Maxillary palp formula (1,2)-4-3-5, segment 4 only slightly longer than segments 1 and 2, segment 3 double long then segment 4, segment 5 almost triple longer than all the others together. Tibial spurs 044, dorsoapical projection from foretibia armed with 4–5 spines. Cuticular structure of sternum V glands forming a pair of elongated rounded ridge. Male genitalia. Segment IX short, its dorsum triangularly slightly roofing over segment X; intersegmental profile between segments IX and X high obtuse angled; stout setae present on apical rim. Segment X elongated low in lateral view rounded digitiform lobes in dorsal view; cerci located middle as a pair of setose warts; vestigial paraproct developed as a pair of ventrolateral straps directed downward from apical lobes. Coxopodite and harpago fused, slightly S-forming and constricted middle. Phallic organ with well visible phallotremal sclerites.

13 14

Centromacronema talan sp. n., holotype, male genitalia: 11 = lateral view; 12 = left gonopod, ventral; 13 = dorsal; 14 = phallus lateral; 15 = phallus apex, ventral

Material examined – Holotype, male: : Huanuco, stream at Carpish, 2500 m, 76°09’W, 9°40’S, 28.VI–3.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (NRM). Paratypes: same data as holotype (2 males, NRM, 2 males, OPC). Chontachaca, Kosnipata-

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)221

Cusco, humid subtropical forest, 700 m, 71°28’03’’W, 13°01’25’’S, 15–30.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (9 males, NRM, 6 males, 1 female, OPC). Huanuco, stream at Chinchao, 2200 m, 76°00’W, 9°66’S, 18–26.VI.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (4 males, 1 female, NRM).

Etymology – Talan, from “-talan”, without in Hungarian, refers to the lost crossvein m resultinginopenmediancell.

Leptonema GUÉRIN-MÉNEVILLE, 1843

Remarks – Here we summarise our understanding of the appendicular and functional genital terminology as applied for this hydropsychid genus (OLÁH &JOHANSON 2008). We follow SCHMID’s (1968, 1970) interpretation suggested for the complex of segments X and XI. We use the term of segment X for this combined complex of segments IX and X. In the Hydropsychidae, segment X either membranous, highly reduced or fused with segment XI. Borders between the segments X and XI are poorly visible in Leptonema, although Y- or variously shaped sutures are present occasionally. Segment XI is represented by cercus and paraproct and associated with periphery of segment X and always located together or nearby. Cerci of podite origin always covered with numerous sensillae and the segmental plates of paraproct having somite origin may have setose sensory, as well as less setose stimulatory or setaless brace elements. Up to three pairs of setose warts are present on the combined segments X and XI highly varying in size, shape, reduction, duplication and fragmentation. FLINT et al. (1987) homologised the wart a, located in a sublateral, dorsal position near the anterior margin of segment X, with the cerci (superior appendages of NIELSEN (1957) or preanal appendages of SCHMID (1968, 1970). Here we treat all the three warts (a, b, c), frequently petiolate and located along the apical margin of mesal sinus, as setose paraproctal processes (dorsoapical and ventroapical setose lobes in several hydropsychid genera) and homologise the setose area present permanently on the lateral lobes with the cerci (superior or preanal appendages). These warts and the ventrolateral sclerotised straps constitute the paraproct. The variously developed and sclerotised ventrolateral straps of the paraproct function like phallic guides forming dorsolateral sheath of the phallocrypt or fused membranously to the basis or dorsum of the phallotheca serving a supplementary role during intromitting movements of the phallic apparatus. The apical section of the phallic organ is varying from simple to very complex assem- blage of membranous or sclerotised lobes, hooks, spines, forks, points patterned by pimp- les, wrinkles, serrated ridges, teeth, plumose, tomentose or tomentulose structures. These structures were symbolised by 9 letters without indicating any homology (FLINT et al. 1987). Here we understand processes a and j as dorsoapical phallothecal tips, process g as ventroapical phallothecal tip and process f as subapical phallothecal outgrowth; process b, b1, and c as apical endothecal sclerites; and paired processes of d and e as subapical endothecal processes.

Annls hist.-nat. Mus. natn. hung. 104, 2012 222 J. Oláh & K. A. Johanson

Leptonema album MOSELY, 1933

Material examined – : Tinalandia Nature Reserve, West Andeal Slope, near Quito, 600 m, 24.XI.2007, light, leg. J. OLÁH Jr. (1 male, OPC).

Leptonema boliviense boliviense MOSELY, 1933

Material examined – : Jujuy, PN, Calilegua, 13 km NW, visitor center, RP 83, 1263 m, 23°41.923’S, 64°52.342’W, 14.I.2008, leg. K. HILL,D.MARSHALL &M. MOULDS (1 male, NRM). : Dept. Pasco, Yanachaga Chemilien NP, INRENA Refugio El Cedro, S10°32.717, W75°21.492, 2460 m, 30.I.2003, leg A. KUN &B.BENEDEK, HNHM expedition (7 males, HNHM).

sp. n. (Figs 16–20)

Diagnosis – This species belongs to the L. amazonense species group, and is similar to L. neblinense FLINT,MCALPINE et ROSS, 1987 from Venezuela. It differs by having intersegmental profile filled, not deep and right-angled; cerci well-developed on the basal region of the lateral lobes, not lacking; mesal lobes of segment X digitate and clavate, not lobe- shaped; phallic apex much dilated and double-winged in ventral view, not simple clavate.

Description – Male. Forewing brown, almost uniformly punctuate with pale spots, forewing length 9 mm. Maxillary palp formula 1-4-(2,3)-5, fourth segment much shorter than third, almost as short as segment first. Tibial spurs 144. Median suture on second abdominal sternum present. Cuticular structure of sternum V glands small-sized. Male genitalia. Segment IX short without discernible pleurosternal lines of secondary longitudinal sutures; and without discernible dorsal keel; intersegmental profile minimal; stout setae present middle on posterior margin. Complex of segment X very specially con- structed by a digitate/clavate mesal lobe and usual lateral lobes, cerci present as well-developed setose warts on basal region of lateral lobes; paraproct developed as a pair of ventrolateral straps directed downward from apical lobes, functioning as sclerotised phallic guides above phallic organ as well as attached to dorsal region of phallic apparatus. Basal segment of gonopod stout, its mesal surface fringed with short stout modified setae. Phallic organ simple with double-winged apex in ventral view; semimembranous articulation of paraproctal lateral strips on basodorsal region of phallotheca for intromitting function and sclerotic basal plate articulation for holding function boths are well visible.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)223 males, OPC). Approuaguekaw, Kaw Mt, 216 mao, 4°33.257’N, 52°11.920’W, Malaise trap, 4–12.II.2007, FRG MF 2, leg. N. JÖNSSON (5 males, NRM, 3 males, OPC).

Etymology – Bunkok, from “bunkók”, clavus plural in Hungarian, refers to the clavate apex of mesal lobes on segment X.

20 17

19 18 16

Leptonema bunkok sp. n., holotype, male genitalia: 16 = lateral view; 17 = left gonopod, ventral; 18 = dorsal; 19 = phallus lateral; 20 = phallus apex, ventral

sp. n. (Figs 21–25)

Diagnosis – This species belongs to the L. pallidum species group, and is similar to L. ramosum FLINT,MCALPINE et ROSS, 1987 from Venezuela, from which it differs by having dorsal keel bluntly triangular, not sharply triangular; intersegmental profile right-angled, not sharp-angled; cerci shifted apicolaterad, not located subapicad; paraproct with 3 small setose lobes, not with 2; endothecal processes on phallic organ differently shaped.

Description – Male. Pale hyaline, probably greenish in life; forewing length 15 mm. Maxillary palp formula (1,4)-3-2-5, fourth segment much shorter than third. Tibial spurs 244. Median suture on second abdominal sternum present, long. Cuticular structure of sternum V glands medium-sized. Male genitalia. Segment IX short with short acrotergit; well developed pleurosternal lines of secondary longitudinal sutures present on lateroventral position above gonopods; triangular dorsal keel short and blunt; intersegmental profile between segments IX and X almost regular right-angled; stout setae present on slope below keel between segments IX

Annls hist.-nat. Mus. natn. hung. 104, 2012 224 J. Oláh & K. A. Johanson and X. Segment X short, constricted laterad by sinus in dorsal view; cerci shifted to apicolaterad; paraproct present as 3 pairs of small setose lobes (warts) apicad of cerci as well as developed as a pair of ventrolateral straps directed downward from apical lobes, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopod, coxopodite with an area of dense setae on inner surface near apex. Phallic organ complex with various endothecal processes.

Material examined – Holotype, male: : Minca, 650 m, 24.III.2011, light, leg. J. OLÁH Jr. (OPC). Paratype: same data as holotype (1 male, OPC).

Etymology – Enikolah dedicated to the daughter of the collector, to ENIKÕ JÚLIA OLÁH.

21 22

24 23

Leptonema enikolah sp. n., holotype, male genitalia: 21 = lateral view; 22 = left gonopod, ventral; 23 = dorsal; 24 = phallus lateral; 25 = phallus apex, ventral

Leptonema harpagum FLINT, MCALPINE et ROSS, 1987

Material examined – San Martin Prov., La Catarata de Ahuashiyascu, 6°27.544’ S, 76°18.192’W, light, loc. 07, 7.I.2009, leg. K. A. JOHANSON & T. MALM (1 male, NRM).

Leptonema inca MOSELY, 1933

Material examined – : River Salvation, Mt Manu, Mother of Good, 546 m, humid subtropical forest, 71°21’22”W, 12°50’16”S, 26–28.II.2005, Malaise trap, leg. JUAN CHAVEZ LOPEZ (5 females, NRM, 1 male, 2 females, OPC).

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)225

Leptonema inspiratum FLINT,MCALPINE et ROSS, 1987

Material examined – : Palotoa, Manu-Mother of Good, 71°18’W, 12°35’S, 436 m, 15–30.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (1 male, NRM).

Leptonema intermedium MOSELY, 1933

Material examined – : Tinalandia Nature Reserve, West Andeal Slope, near Quito, 600 m, 24.XI.2007, light, leg. J. OLÁH Jr. (3 males, OPC). Tinalandia Nature Reser- ve, West Andeal Slope, near Quito, 600 m, 15.X.2010, light, leg. J. OLÁH Jr. (3 males, OPC). Tinalandia Nature Reserve, West Andeal Slope, near Quito, 600 m, 19.XIII.2011, light, leg. J. OLÁH Jr. (2 males, OPC).

sp. n. (Figs 26–31)

Diagnosis – This species belongs to the L. stigmosum species group, and is most similar to L. spirillum FLINT,MCALPINE et ROSS, 1987, from Peru, but differs by having the head of phallic organ differently shaped.

Description – Male. Pale ochraceous hyaline, probably greenish in life; forewing length 17 mm. Maxillary palp formula 4-1-3-2-5, fourth segment much shorter than third. Tibial spurs 244. Median suture on second abdominal sternum present, long. Cuticular structure of sternum V glands small. Male genitalia. Segment IX short; well developed pleurosternal lines of secondary longitudinal sutures present on lateroventral position above gonopods; dorsal keel rounded blunt; intersegmental profile between segments IX and X right-angled; stout setae present on tip of keel and on posterior ridge of segments IX. Segment X short, constricted basad and excised mesad in dorsal view; cerci shifted to apicolaterad; paraproct composed of a single pair of short apical warts (damaged on left side!) and a pair of ventrolateral straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopod, coxopodite without an area of dense setae on inner surface near apex. Phallic organ simple with 2 pairs of feathered endothecal processes; combined b and c present as apical pair of anterad recurving and downward directed very stout processes with spinose head and spinose process e; process g, ventroapical apex of phallotheca modified into a narrowing median lobe, laterad serrated process a; dorsoapical head of phallotheca bilobed; phallotremal sclerites dark pigmented triangular.

Material examined– Holotype, male: : Western Andean Slope, Alambi, 24.IX.2010, light, leg. J. OLÁH Jr. (OPC). Paratypes: same data as holotype (4 females, OPC).

Etymology – Janolah dedicated to the collector, JÁNOS OLÁH Jr.

Annls hist.-nat. Mus. natn. hung. 104, 2012 226 J. Oláh & K. A. Johanson

27 30

31 28

26 29

Leptonema janolah sp. n., holotype, male genitalia: 26 = right lateral view; 27 = left gonopod, ventral; 28 = dorsal; 29 = phallus lateral; 30 = phallus apex, ventral; 31 = phallus apex, dorsal

Leptonema ketos sp. n. (Figs 32–36)

Diagnosis – This species belongs to the L. amazonense species group, and is similar to L. irroratum FLINT, 1974 from Surinam. Leptonema ketos sp. n. has male genital architec- ture of L. irroratum, but differs by having cerci divided into double warts, not single wart; setose paraproctal warts differently shaped; especially the mesal pair is modified into short irregular processes, not long digitate; double-lobed pattern of phallic apex significantly modified in ventral view.

Description – Male. Forewing brown almost uniformly punctuate with pale spots, forewing length 10 mm. Maxillary palp formula 1-4-(2,3)-5, fourth segment much shorter than third, almost as short as segment first. Tibial spurs 144. Median suture on second abdominal sternum present. Cuticular structure of sternum V glands medium-sized. Male genitalia. Segment IX short without discernible pleurosternal lines of secondary longitudinal sutures; dorsal keel triangular in dorsal view; intersegmental profile filled; 5–6 stout setae present along posterior margin. Complex of segment X composed of a basodorsal rounded setose lobe, a pair of mesal short irregular processes and usual lateral lobes, cerci present as duplicated warts on middle constricted region of lateral lobes; paraproct developed as a pair of ventrolateral straps directed downward from apical lobes; its basal section well developed and separated. Basal segment of gonopod stout supplied with a basomesal short process; harpagones clavate. Phallic organ simple with double-winged apex in ventral view.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)227

Material examined – Holotype, male: : Approuaguekaw, Kaw Mt, 104 mao, 4°33.035’N, 52°11.661’W, Malaise trap, 23.I–7.II.2007, FRG MF3, leg. N. JÖNSSON (NRM). Paratypes: same data as holotype (3 males, NRM). Approuaguekaw, Kaw Mt, 216 mao, 4°33.257’N, 52°11.920’W, Malaise trap, 4–12.II.2007, FRG MF 2, leg. N. JÖNSSON (1 male, NRM, 1 male, OPC).

Etymology – Ketos, from “kettõs”, doubled in Hungarian, refers to the cerci composed of duplicated warts.

Remarks –FLINT et al. (1987) found specimens from Venezuela that agree in general structure in all parts of the male genitalia with those of L. irroratum FLINT, 1974, but almost every part differs slightly in shape or proportionate length.

32 33

35 34

Leptonema ketos sp. n., holotype, male genitalia: 32 = lateral view; 33 = left gonopod, ventral; 34 = dorsal; 35 = phallus lateral; 36 = phallus apex, ventral

sp. n. (Figs 37–41)

Diagnosis – This species belongs to the L. plicatum species group, and is closest to L. andinum FLINT, 1991 from Colombia, from which it differs by having the head of phallic organ differently shaped.

Description – Male. Pale ochraceous hyaline, probably greenish in life; forewing length 217 mm. Maxillary palp formula 4-1-3-2-5, fourth segment much shorter than third. Tibial spurs 244. Median suture on second abdominal sternum present, long. Cuticular structure of sternum V glands small.

Annls hist.-nat. Mus. natn. hung. 104, 2012 228 J. Oláh & K. A. Johanson

Male genitalia. Segment IX short; well developed pleurosternal lines of secondary longitudinal sutures present on lateroventral position above gonopods; dorsal keel rounded triangular; intersegmental profile between segments IX and X indistinct; stout setae present on tip of keel and on posterior ridge of segments IX. Segment X short, constricted basad and excised mesad in dorsal view; cerci shifted to apicolaterad; paraproct composed of warts a and b fragmented along margin of mesal excision and of a pair of internal straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopod, coxopodite without an area of dense setae on inner surface near apex. Phallic organ simple with a single pair of endothecal processes (combined b and c), present as apical pair of anterad recurving spines with lateral larger and mesal smaller arms; process g, ventroapical apex of phallotheca fused, membranous, in- distinctly separated from heavily sclerotised spines of process b and c, dorsoapical head of phallotheca bilobed; phallotremal sclerites dark pigmented triangular; a pair of serrulated dorsal lines present subapicad.

Material examined – Holotype, male: : dept. Pasco, Yanachaga Chemilien NP, INRENA Refugio El Cedro, S10°32.717, W75°21.492, 2460 m, 30.I.2003, leg A. KUN &B. BENEDEK, HNHM expedition (HNHM).

Etymology – Kunbenorum dedicated to the collectors.

37 38

39 40

Leptonema kunbenorum sp. n., holotype, male genitalia: 37 = lateral view; 38 = left gonopod, ventral; 39 = dorsal; 40 = phallus lateral; 41 = phallus apex, ventral

Leptonema maculatum MOSELY, 1933

Material examined – : Approuaguekaw, Kaw Mt., 216 mao, 4°33.257’ N, 52°11.920’W, Malaise trap, 4–12.II.2007, FRG MF 2, leg. N. JÖNSSON (2 males, NRM).

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)229

Approuaguekaw, Kaw Mt., 104 mao, 4°33.035’N, 52°11.661’W, Malaise trap, 23.I–7.II.2007, FRGMF3,leg.N.JÖNSSON (3 males, NRM, 2 males, OPC). Approuaguekaw, Kaw Mt, 4°33.035‘N, 52°11.661‘W, 104 mao 23.I.2007, FRG 4, leg. N. JÖNSSON (2 males, NRM).

sp. n. (Figs 42–45)

Diagnosis – This species belongs to the L. stigmosum species group, and is most similar to L. boliviense MOSELY, 1933 from Bolivia, from which it differs in having intersegmental profile very low and obtusely angled, not sharply angled; the paraproctal setal wart pattern is different; the harpago is capitate, not parallel-sided; and the phallic apex is different.

Description – Male. Pale ochraceous hyaline, probably greenish in life; forewing length 18 mm. Maxillary palp formula 4-1-3-2-5. Tibial spurs 244. Median suture on second abdominal sternum present, long. Cuticular structure of sternum V glands small ovoid.

42 43

44 45

Leptonema magas sp. n., holotype, male genitalia: 42 = lateral view; 43 = left gonopod, ventral; 44 = dorsal; 45 = phallus lateral

Male genitalia. Segment IX short; well-developed pleurosternal lines of secondary longitudinal sutures present on lateroventral position above gonopods; dorsal keel reduced, indistict; intersegmental profile between segments IX and X low and obtuse-angled; stout setae present on dorsal posterior ridge of segments IX. Segment X short, highly constricted basad in dorsal view; membranous median lobes well-developed; lateral lobes produced laterad in dorsal view; cerci spread on larger surface of lateral lobes; paraproct present as warts a, b on inner sinus and a pair of ventrolateral straps directed downward from apical

Annls hist.-nat. Mus. natn. hung. 104, 2012 230 J. Oláh & K. A. Johanson margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopod, coxopodite without an area of dense setae on inner surface near apex, harpagones capitate. Phallic organ simple with 3 pairs of feathered endothecal processes: combined b and c present as apical pair of spinose anterad recurving and downward directed stout processes and less spinose processes d and e; process g, ventroapical apex of phallotheca membranous, indistinct; process a, dorsoapical head of phallotheca bilobed; phallotremal sclerites dark pigmented, triangular.

Material examined – Holotype, male: : San Martin Prov., river crossing rd. Olmos-Tarapoto, 385 km (rd.) E Olmos Desv. Jaén, 5°40.055’S, 77°43.396’W, light, loc. 04, 5.I.2009, leg. K. A. JOHANSON & T. MALM (NRM).

Etymology – Magas, from “magas”, high in Hungarian, refers to the high basal section of phallotheca.

Leptonema mandibulatum FLINT, MCALPINE et ROSS, 1987

Material examined – : San Martin Prov., Rio Huallaga, at Pumarihri Huallaga Lodge, between Juan Guerra and Chazuta, 14 km (rd) W Chazuta, 6°36.643’S, 76°12.555’W, light, loc. 10, 9.I.2009, leg. K. A. JOHANSON &T.MALM (4 males, 1 female, NRM, 3 males, 1 female, OPC). San Martin Prov., stream crossing Juan Guerra-Chazuta rd., 10 km (rd.) W Chazuta, 6°37.157’S, 76°10.905’W, light, loc. 12, 10.I.2009, leg. K. A. JOHANSON &T. MALM (1 male, NRM). San Martin Prov., creek crossing rd. Juan Guerra-Chazuta, 14 km (rd.) E Colombia Bridge, 6°35.594’S, 76°13.172’W, light, loc. 09, 9.I.2009, leg. K. A. JOHANSON & T. MALM (1 male, NRM).

sp. n. (Figs 46–50)

Diagnosis – This species belongs to the L. complexum species group, and is closest to L. inca MOSELY, 1933 from Peru. The pattern of the male genitalia is similar, but almost all the genital substructures are differently shaped: pattern of paraproctal setose warts; liguliform lateral projection of paraproctal ventrolateral straps; harpago parallel-sided and capitate, not with mesal hump, that is not indicated on the original drawings of MOSELY (1933); all the processes are differently shaped on the phallic head.

Description – Male. Pale hyaline, probably greenish in life; forewing length 15 mm. Maxillary palp formula 1-4-3-2-5. Tibial spurs 244. Median suture on second abdominal sternum present, long. Cuticular structure of sternum V glands large ovoid. Male genitalia. Segment IX short; well developed pleurosternal lines of secondary longitudinal sutures present on lateroventral position above gonopods; dorsal keel very narrow digitiform with an apical less sclerotised blunt process; intersegmental profile

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)231 between segments IX and X right-angled; stout setae present on dorsal posterior region of segments IX. Segment X short; cerci spread on larger surface of lateral lobes; setose components of paraproct present on inner sinus as a pair of basomesal compact warts and apical diffused pair of setose fringed lobes (warts a, b); a pair of ventrolateral straps directed downward from apical margin, and produced liguliform lateral lobes with setose apex. Basal segment of gonopod, coxopodite without an area of dense setae on inner surface near apex; harpagones parallel-sided and capitate. Phallic organ with all processes present, except j and g and characterized by strongly developed platelike unpaired single process f; phallotremal sclerites dark pigmented triangular.

Material examined – Holotype, male: : San Martin Prov., La Catarata de Ahua- shiyascu, 6°27.544’S, 76°18.192’W, light, loc. 07, 7.I.2009, leg. K. A. JOHANSON &T.MALM (NRM). Paratypes: same data as holotype (5 males, NRM, 3 males, OPC).

Etymology – Masinca, from “más inca”, different inca in Hungarian, refers to the well differing, but close relation to L. inca.

46 47

48 49

Leptonema masinca sp. n., holotype, male genitalia: 46 = lateral view; 47 = left gonopod, ventral; 48 = dorsal; 49 = phallus lateral; 50 = phallus apex, ventral

Leptonema mastigion FLINT, MCALPINE et ROSS, 1987 (Figs 51–55)

Material examined – : Tinalandia Nature Reserve, West Andean Slope, near Quito, 600 m, 19.XIII.2011, light, leg. J. OLÁH Jr. (2 males, OPC).

Annls hist.-nat. Mus. natn. hung. 104, 2012 232 J. Oláh & K. A. Johanson

51 52

Leptonema mastigion FLINT,MCALPINE et ROSS, 1987, male genitalia: 51 = lateral view; 52 = left gonopod, ventral; 53 = dorsal; 54 = phallus lateral; 55 = phallus apex, ventral

sp. n. (Figs 56–60)

Diagnosis – This species belongs to the L. complexum species group, and is most similar to L. inca MOSELY, 1933 and L. masinca sp. n., both from Peru. The new species is distinguished from both by having microtrichia covered dorsal keel of segment IX triangular, not digitiform in dorsal view; irregular apical warts of paraproct (b) not longer than lateral lobes; harpagones distinctly capitate; processes on phallic head differently formed.

Description – Male. Pale hyaline, probably greenish in life; forewing length 14 mm. Maxillary palp formula 1-4-3-2-5. Tibial spurs 244. Median suture on second abdominal sternum present, long. Cuticular structure of sternum V glands large ovoid. Male genitalia. Segment IX short; well developed pleurosternal lines of secondary longitudinal sutures present on lateroventral position above gonopods; dorsal keel triangular with an apical less sclerotised blunt process; intersegmental profile between segments IX and X right-angled; stout setae present on dorsal posterior region of segments IX. Segment X short; cerci spread on larger surface of lateral lobes; setose components of paraproct present on inner sinus as a pair of basomesal compact warts (wart a) and apical diffused pair of setose fringed lobes (wart b); a pair of ventrolateral straps directed downward from apical margin, and produced liguliform lateral lobes with setose apex. Basal segment of gonopod, coxopodite without an area of dense setae on inner surface near apex; harpagones parallel-sided and distinctly capitate. Phallic organ with all processes present, except j and g and characterized by strongly developed bifid apex of fused process b; phallotremal sclerites dark pigmented triangular.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)233

Material examined – Holotype, male: : San Martin Prov., creek crossing rd. Juan Guerra-Chazuta, 14 km (rd.) E Colombia Bridge, 6°35.594’S, 76°13.172’W, light, loc. 09, 9.I.2009, leg. K. A. JOHANSON &T.MALM (NRM). Paratypes: same data as holotype (1 male, NRM, 1 male, OPC).

Etymology – Meginca, from “még inca”, another or more inca in Hungarian, refers to again a new species close to L. inca.

56 57

58 59

Leptonema meginca sp. n., holotype, male genitalia: 56 = lateral view; 57 = left gonopod, ventral; 58 = dorsal; 59 = phallus lateral; 60 = phallus apex, ventral

sp. n. (Figs 61–65)

Diagnosis – This species belongs to the L. stigmosum species group, and is most similar to L. neadelphus FLINT,MCALPINE et ROSS, 1987 from Colombia, but differs by having the paraproctal warts and the head of phallic organ differently structured.

Description – Male. Pale ochraceous hyaline, probably greenish in life; forewing length 21 mm. Maxillary palp formula 1-4-3-2-5. Tibial spurs 244. Median suture on second abdominal sternum present, long. Cuticular structure of sternum V glands small ovoid. Male genitalia. Segment IX short; well developed pleurosternal lines of secondary longitudinal sutures present on lateroventral position above gonopods; dorsal keel triangular with an apical less sclerotised sharp triangular continuation; intersegmental profile between segments IX and X right-angled; stout setae present on dorsal posterior ridge of segments IX. Segment X short; cerci spread on larger surface of lateral lobes; paraproct present on inner sinus as a basomesal compact and apical diffused pair of setose lobes (warts a, b)

Annls hist.-nat. Mus. natn. hung. 104, 2012 234 J. Oláh & K. A. Johanson and a pair of ventrolateral straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopod, coxopodite without an area of dense setae on inner surface near apex. Phallic organ simple with 2 pairs of feathered endothecal processes: combined b and c present as apical pair of strongly spinose anterad recurving and downward directed stout processes and spinose processes e; process g, ventroapical apex of phallotheca membranous, indistinct; process a, dorsoapical head of phallotheca bilobed; phallotremal sclerites dark pigmented triangular.

Material examined – Holotype, male: : Amazonas Prov., river crossing Olmos- Tarapoto rd., 71 km (rd.) E Olmos Desv. Jaén, 5°41.178’S, 77°46.421’W, light, loc. 02, 5.I. 2009, leg. K. A. JOHANSON &T.MALM (NRM). Paratypes: same data as holotype (6 males, NRM, 4 males, OPC).

Etymology – Named for the type locality.

62 61

65 63

Leptonema olmos sp. n., holotype, male genitalia: 61 = lateral view; 62 = left gonopod, ventral; 63 = dorsal; 64 = phallus lateral; 65 = phallus apex, ventral

Leptonema pallidum GUERIN, 1843

Material examined – : Approuaguekaw, Kaw Mt, 104 mao, 4°33.035’ N, 52°11.661’W, Malaise trap, 23.I–7.II.2007, FRG MF3 leg. N. JÖNSSON (2 females, OPC; 1 male, 1 female, NRM).

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)235

Leptonema plicatum MOSELY, 1933

Material examined – : State of Veracruz, Los Tuxtlas, around the Estacion the Biologia Los Tuxtlas, N 18°35.213’, W 095°04.462’, 30 mao, 27.VI.2006, light trap, leg. M. ESPELAND & T. MALM (2 males, 2 females, NRM, 1 male, 1 female, OPC).

Leptonema rosenbergi MOSELY, 1933

Material examined – : Lita, IX.2008, light, leg. J. OLÁH Jr. (1 male, OPC).

Leptonema sparsum (ULMER, 1905)

Material examined – : San Martin Prov., Rio Negro, 37 km (rd.) W Moyobamba, near Olmos-Tarapoto rd., 6°00.278’S, 77°15.437’W, light, loc. 05, 6.I.2009, leg. K. A. JOHANSON & T. MALM (1 male, NRM).

Leptonema spirillum FLINT, MCALPINE et ROSS, 1987

Material examined – : Otun Quinbaya National Park, La Suiza, 1900 m, 13.III.2011, leg. J. OLÁH Jr. (1 male, OPC). : Huanuco, stream at Chinchao, 2200 m, 76°00’W, 9°66’S, 18–26.VI.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (1 male, 14 females, NRM). Huanuco, stream at Carpish, 2500 m, 76°09’W, 9°40’S, 28.VI–3.VII. 2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (1 male, 13 females, OPC). Dept. Pasco, Yanachaga Chemilien NP, INRENA Refugio El Cedro, S10°32.717, W75°21.492, 2460 m, 30.I.2003, leg A. KUN &B.BENEDEK, HNHM expedition (4 males, HNHM). Pasco Reg., Yanachaga-Chemillen NP, side river to Rio Huancabamba, N end of park, along Oxabamba- Pozuzo rd., 10°11.133’S, 75°34.106’W, light, loc.01.31.XII.2008, leg. K. A. JOHANSON &T. MALM (9 males, 1 female, NRM).

sp. n. (Figs 66–70)

Diagnosis – This species belongs to the L. pallidum group, and is closest to L. viridianum NAVÁS, 1916 from Brazil, from which it differs by having paraproctal setose warts differently shaped; ventrobasal lobe of gonocoxite short and blunt, not long digitiform; phallothecal and endothecal processes on phallic apex different, processes b plumose, not branched.

Description – Male. Pale hyaline, probably greenish in life; forewing length 12 mm. Maxillary palp formula 4-1-3-2-5, fourth segment much shorter than third. Tibial spurs 244.

Annls hist.-nat. Mus. natn. hung. 104, 2012 236 J. Oláh & K. A. Johanson

Median suture on second abdominal sternum present, long. Cuticular structure of sternum V glands medium-sized. Male genitalia. Segment IX short with short acrotergit; well developed pleurosternal lines of secondary longitudinal sutures present on lateroventral position above gonopods; sharp triangular dorsal keel long; intersegmental profile between segments IX and X obtuse angled; stout setae present on slope below keel between segments IX and X. Segment X short; cerci long on lateral lobes; paraproctal setose warts present as 23 pairs of small setose lobes (warts); a pair of paraproctal ventrolateral straps directed downward from apical lobes, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopod, coxopodite without distinct area of dense setae on inner surface near apex. Phallic organ complex with various endothecal processes, process b plumose.

Etymology – Tollas, from “tollas”, feathered in Hungarian, refers to the plumose endothecal process b on phallic apex.

66 67

69 68

Leptonema tollas sp. n., holotype, male genitalia: 66 = lateral view; 67 = left gonopod, ventral; 68 = dorsal; 69 = phallus lateral; 70 = phallus apex, ventral

Leptonema trifidum FLINT, MCALPINE et ROSS, 1987

Material examined – : Chontachaca, Kosnipata-Cusco, humid subtropical forest, 700 m, 71°28’03”W, 13°01’25”S, 15–30.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (10 males, NRM, 5 males, 1 female, OPC). Palotoa, Manu-Mother of Good, 71°18’W, 12°35’S, 436 m, 15–30.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (8 males NRM, 4 males, OPC). Dept. Pasco, Yanachaga Chemilien NP, INRENA Refugio El Cedro, S10°32.717, W75°21.492, 2460 m, 30.I.2003, leg A. KUN &B.BENEDEK,

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)237

HNHM expedition (1 male, HNHM). San Martin Prov., La Catarata de Ahuashiyascu, 6°27.544’S, 76°18.192’W, light, loc. 07, 7.I.2009, leg. K. A. JOHANSON &T.MALM (3 males, NRM). San Martin Prov., creek crossing rd. Tarapoto-Yurimaguas, ca. 30 km (rd.) NE Tarapoto, 6°24.904’S, 76°18.756’W, light, loc. 08, 8.I.2009, leg. K. A. JOHANSON &T. MALM (3 males, 3 females, NRM).

Leptonema viridianum NAVÁS, 1916

Material examined – : Gareno, near Puerto Napo, 10–11.X.2010, light, leg. J. OLÁH Jr. (1 male, 1 female, OPC). : Palotoa, Manu-Mother of Good, 71°18’W, 12°35’S, 436 m, 15–30.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (1 male, NRM, 2 males, OPC). San Martin Prov., Rio Huallaga tributary, small river passing Cha- zuta, 6°34.665’S, 76°08.209’W, light, loc. 11, 10.I.2009, leg. K. A. JOHANSON &T.MALM (2 males, NRM).

Macronema PICTET, 1836 Macronema bifidum FLINT, 1974

Material examined – : Approuaguekaw, Kaw Mt, 104 m, 4°33.035’N, 52°11.661’W, Malaise trap, 4–12.II.2007, FRG MF3, leg. N. JÖNSSON (1 male, NRM).

Remarks – The horizontal section of phallotheca, the stem forming a regular parallel- sided downcurving tube, not irregular in lateral view; paraproctal apices blunt, not pointed in lateral view. Variability study is required to separate as this specimen from French Guiana as a new species.

Macronema burmeisteri BANKS, 1924

Material examined – : Manu-Mother of Good, River Tonchaca, 546 m, 71°20’ W, 12°50’S, 436 m, 17–22.V.2005,Malaise and light trap, leg. FAVIOLA MONTES CARLOS & JUAN CHAVEZ (1 female, NRM).

Macronema fragile BANKS, 1915

Material examined – : Approuaguekaw, Kaw Mt, 104 mao, 4°33.035’ N, 52°11.661’W, Malaise trap, 23.I–7.II.2007, FRG MF3, leg. N. JÖNSSON (2 males, NRM). Approuaguekaw, Kaw Mt, 216 mao, 4°33.257’N, 52°11.920’W, Malaise trap, 4–12.II.2007, FRG MF 2, leg. N. JÖNSSON (2 males, 2 females, NRM, 1 male, 2 females, OPC).

Annls hist.-nat. Mus. natn. hung. 104, 2012 238 J. Oláh & K. A. Johanson

Macronema fraternum BANKS, 1910

Material examined – : Approuaguekaw, Kaw Mtn, 104 m, 4°33.035’ N, 52°11.661’W, Malaise trap, 23.I–7.II.2007,FRG MF3, leg. N. JÖNSSON (1 male, NRM).

sp. n. (Figs 71–75)

Diagnosis – This species is close to M. fraternum BANKS, 1910 from Guiana, from which it differs by having forewings with entirely brown unicolour membrane except pale transversal band on anastomosis; the internal pattern of the phallic apex is different; and the dorsal tip of the phallotheca is bilobed, not monolobed.

Description – Male. Dorsum dark brown, ventrum and appendages paler; forewing membrane brown with pale transversal band on anastomosal area; forewing length 8.2 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 044. Cuticular structure of sternum V glands forming elongated and rounded ridge. Male genitalia. Segment IX very short, dorsal keel forming a bilobed tip; intersegmental profile between segments IX and X obtuse angled. Segment X triangular in lateral view with widely streaching lateral lobes in dorsal view; cerci located middle as a pair of small setose warts; paraproct present as more sclerotised ventrolateral straps, functioning as sclerotised phallic guides encircling above and almost below phallic organ. Gonopods undivided. Dorsal tip of phallotheca bilobed.

71 72

74 73

Macronema ketleben sp. n., holotype, male genitalia: 71 = lateral view; 72 = left gonopod, ventral; 73 = dorsal; 74 = phallus lateral; 75 = phallus apex, ventral

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)239

Material examined – Holotype, male: : Approuaguekaw, Kaw Mt, 216 m, 4°33.257’N, 52°11.920’W, Malaise trap, 19.I–4.II.2007, FRG MF 2, leg. N. JÖNSSON (NRM). Paratype: Approuaguekaw, Kaw Mt, 104 m, 4°33.035’N, 52°11.661’W, Malaise trap, 4–12.II.2007, FRG MF3, leg. N. JÖNSSON (1 male, OPC).

Etymology – Ketleben, coined from “kétlebeny”, two-lobed in Hungarian, refers to the mesaly excised dorsal tip of phallotheca that forms in the bilobed terminal structure.

Remarks – The genital structure of the paratype has some modifications: dorsal keel monolobed, not bilobed; horizontal section of phallotheca strongly curved, not only slightly. More specimens are required to examine the variability of these characters, in order to separate the paratype as a new species.

sp. n. (Figs 76–79)

Diagnosis – This species is similar to M. bifidum FLINT, 1974 from Surinam, from which it differs by having forewings with a light irregular transverse membrane band on a- nastomosis; the dorsum of segment IX is elevated vertically in lateral view, not horizon- tally; the ventrum has double pair of processes, not with single; the paraproct is oriented more vertically, not obliquely horizontal; lateral profile of phallotehca different.

Description – Male. Dorsum dark brown, ventrum and appendages paler; forewing membrane brown with pale transversal band on anastomosal area; forewing length 11 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 044. Cuticular structure of sternum V glands forming elongated and rounded ridge without any filament. Male genitalia. Segment IX very short, dorsal keel forming a vertically elevated, monolobed tip; ventrum with a pointed lateral and an obtuse mesal pair of processes; intersegmental profile between segments IX and X obtuse angled. Segment X triangular in lateral view with widely streaching lateral lobes in dorsal view; cerci located middle as a pair of short digitate setose warts; paraproct present as more sclerotised ventrolateral straps, functioning as sclerotised phallic guides encircling above and almost below phallic organ. Gonopods undivided. Dorsal tip of phallotheca monolobed, lateral surface sorrounding ventroapical opening densely speculate; internal sclerotic complex indistinct.

Material examined – Holotype, male: : Approuaguekaw, Kaw Mt, 104 mao, 4°33.035’N, 52°11.661’W, Malaise trap, 23.I–7.II.2007, FRG MF3, leg. N. JÖNSSON (1 male, NRM). Paratype: same data as holotype (4 males, NRM, 1 male, OPC). Approuaguekaw, Kaw Mt, 104 m, 4°33.035’N, 52°11.661’W, Malaise trap, 4–12.II.2007, FRG MF3, leg. N. JÖNSSON (2 males, NRM, 2 males OPC).

Etymology – Toblet, coined from “többlet”, surplus in Hungarian, refers to the lateral pointed processes on the ventral margin of segment IX surplus to the mesal pair of blunt processes present on ventrum IX.

Annls hist.-nat. Mus. natn. hung. 104, 2012 240 J. Oláh & K. A. Johanson

77 78 79

Macronema toblet sp. n., holotype, male genitalia: 76 = lateral view; 77 = left gonopod, ventral; 78 = dorsal; 79 = phallus lateral

Macrostemum KOLENATI, 1859 Macrostemum erichsoni (BANKS, 1920)

Material examined – : Approuaguekaw, Kaw Mt, 104 m, 4°33.035’N, 52°11.661’W, Malaise trap, 4–12.II.2007, FRG MF3, leg. N. JÖNSSON (1 female, NRM).

sp. n. (Figs 80–84)

Diagnosis – This species is most similar to M. hyalinum (PICTET, 1836) from Brazil, Colombia, Guyana, Peru and Venezuela. It differs by having a more faded forewing pattern, not brightly contrasted; the 2 complete middle transversal light bands running from anterior to posterior margins, not fragmented; segment X pointed, not rounded in lateral view; dorsal excision is widely U-shaped, not narrow V-shaped; the harpagones are longer than half length of the coxopodites, not shorter; the lateral profile of the phallic apex is upward rounded, not straight.

Description – Male (in alcohol). Pale hyaline, probably greenish in life; forewing length 12 mm. Head, thoracic sclerites and forleg femur dark castanean brown; antennae with sacpus, palpi, haustellum, warts and legs yellowish. Maxillary palp formula 1-2-4-3-5, first 4 segments almost subequal. Tibial spurs 244. Cuticular structure of sternum V glands forming a long filament.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)241

Male genitalia. Segment IX short, ventrum only little longer; dorsolateral and ventrolateral lobes with stout setae short; small triangular dorsal keel short; intersegmental profile between segments IX and X obtuse angled. Segment X short, pointed apicad in lateral view, median excision wide U-shaped; cerci shifted apicad as setose surface; paraproct present as more sclerotised ventrolateral straps, functioning as sclerotised phallic guides encircling above phallic organ. Harpagones longer than half length of coxopodite. Lateral profile of phallic apex rounding upwards.

Material examined – Holotype, male: : San Martin Prov., La Catarata de Ahua- shiyascu, 6°27.544’S, 76°18.192’W, light, loc. 07, 7.I.2009, leg. K. A. JOHANSON &T.MALM (1 male, NRM). Paratype: same data as holotype (1 male, OPC).

Etymology – Felker, coined from “felfelé kerekedõ”, rounding upward in Hungarian, refers to the lateral profile of phallic apex.

83 81 84

Macronema felker sp. n., holotype, male, right forewing and genitalia: 80 = right forewing; 81 = lateral view; 82 = left gonopod, ventral; 83 = dorsal; 84 = phallus lateral

Macrostemum hyalinum (PICTET, 1836)

Material examined – : Chontachaca, Kosnipata-Cusco, humid subtropical forest, 700 m, 71°28’03”W, 13°01’25”S, 15–30.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS. (2 females, NRM, 1 male, 1 female, OPC). River Salvation, Mt Manu, Mother of Good, 546 m, humid subtropical forest, 71°21’22”W, 12°50’16”S, 26–28.II. 2005, Malaise trap, leg. JUAN CHAVEZ LOPEZ (5 males, NRM, 2 males, 1 female, OPC).

Annls hist.-nat. Mus. natn. hung. 104, 2012 242 J. Oláh & K. A. Johanson

Macrostemum ulmeri (BANKS, 1913)

Material examined – : Huanuco, stream at Carpish, 2500 m, 76°09’W, 9°40’S, 28.VI–3.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (1 female, NRM). Chonta- chaca, Kosnipata-Cusco, humid subtropical forest, 700 m, 71°28’03’’W, 13°01’25”S, 15–30. VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (3 males, 1 female, NRM, 2 males, 1 female, OPC). Palotoa, Manu-Mother of Good, 71°18’W, 12°35’S, 436 m, 15–30.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (1 male, 2 females, NRM). Manu-Mother of Good, River Tonchaca, 546 m, 71°20’W, 12°50’S, 436 m, 17–22.V.2005, Malaise and light trap, leg. FAVIOLA MONTES CARLOS &JUAN CHAVEZ (1 male, NRM).

Plectromacronema ULMER, 1906 Plectromacronema comptum ULMER, 1906

Material examined – : St-Laurentdu-Maroni: Maroni River: A- Chimato-Campop, village 5°13.716’N, 54°17.498’W, 0 m, 6.II.2007, FRG 12, leg. N. JÖNSSON (1 male, 3 females NRM, 1 male, 3 female, OPC).

Smicridea MCLACHLAN, 1871

Remarks – A very diverse and abundant genus in the Neotropical realm distributed as far north as the southwestern United States, and with close relatives in Australia. More studies on species grouping is highly required. Here we summarise our present state of know- ledge on species groups (Table 1).

Smicridea (Smicridea) annulicornis (BLANCHARD, 1851)

Material examined – : Region del Biobio (VIII), ca. 20 km WSW Arauco, stream crossing road at Puento Caripilum, 37°18.141’S, 73°30.633’W, 20 m (Loc #06), light trap 2.I.2006, K. A. JOHANSON (19 males, 14 females, NRM). Region de los Lagos (X), Isla Gran- de de Chiloé, Rio Melilebú, along road between Tebuhueico and Hullinco, 5.4 km S crosspoint to Curaco, river, 100 m upstream bridge, ca. 42°43.089’S, 73°53.797’W, 43 m (Loc #15), net, leg. K.A.JOHANSON (1 male, NRM). Region de los Lagos (X), PN, Vicente Rosales, 200 m downstream Saltos de Petrohué, 6 km SW Petrohué and Lago Todo los Santos, stream parallel to Rio Petrohué, 41°10.378’S, 72°27.057’W, 110 m (Loc #18), light trap 12.I.2006, K. A. JOHANSON (7 males, NRM, 4 males, OPC). Region de la Araucania (IX), Rio Tilquilco, 50mdownstream bridge at 1.2kmS entrance to PN, Huerquehue 39°10.693’S,71°44.269’W, 776 m (Loc #21), light trap 13.I.2006, K. A. JOHANSON (12 males, OPC). Region del Maule (VII), river ca. 43.4 km ESE Curicó, Potrero Grande, at Puente los Morongos, 35°12.990’S, 70°58.517’W, 727 m (Loc #31), light trap 20.I.2006, K. A. JOHANSON (9 males, NRM, 9 males, OPC).

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)243

Main features of examined species in species groups and complexes of Smicridea (Smicridea) and Smicridea (Rhyacophylax) subgenera based mostly on FLINT (1974)

Species groups Characters forewing posterior crossveins (m-cu, cu) far anteriad from anterior crossveins (m, r-m, r); hindwing radiomedial veins well sepa- raterd from Cu; cuticular structure of sternum V glands with short anterolateral filament; male with 2 pairs of internal, reticulate sacs of pheromonal glands in abdominal segments VI and VII fasciatella group phallotheca simple, angulate tube with spines and internal sclerites at apex nigripennis group phallotheca complex, open posterodorsally with spines and processes

forewing crossveins are nearly in alignment; hindwing radiomedial veins closely approximate to Cu; cuticular structure of sternum V glands with long anterolateral filament; without internal, reticulate abdominal sacs argentina group phallotheca with 2-3 small dark apical points argentina complex phallotheca slender in lateral view: argentina, homora sp. n., palmar discalis complex phallotheca robust in lateral view: discalis, magdalena, scutellaria dispar group phallotheca simple without major apical lobes: dispar, dithyra, ve- racruzensis appendiculata complex phallotheca withou major apical lobes, phallotremal sclerites elongated scissorlike: appendiculata, flinti, mangaratiba, roraimense, sarvaka sp. n., voluta murina group pair of hornlike apicodorsal lobes present on apex of phallotheca: murina, zanclophora peruana group phallotheca with asperites on external surface and rather elabora- ted eversible internal complex: acuminata, andicola, biserulata, lebena sp. n., karcsa sp. n., marua, peruana, petasata, probolophora, sarkoska sp. n., spinulosa, sudara sp. n., ventridenticulata, villa sp. n. radula group apex of phallotheca with a pair of radulalike ventrolateral lobes: kapara sp. n., radula, resela sp. n., pseudoradula, vakara sp. n. signata group vestigial paraproct with spiny processes developed in various num- ber and position on ventrolateral margin of segment X; apex of aedeagus with fixed tongue-like ventromesal process: arizonensis, bidactyla, bifurcata, hajla, inarmata, leloga sp. n., lobata, nemtompa, pseudolobata, signata, singri, tavola salta complex single hook-like spiny process midway on paraproct: fogasa, kampoka, nemora, salta

Annls hist.-nat. Mus. natn. hung. 104, 2012 244 J. Oláh & K. A. Johanson

sp. n. (Figs 85–89)

Diagnosis – This species has a phallic organ with unusually low phallobase, similar to S. (R.) hajla sp. n., S. (R.) nemtompa sp. n. and S. (R.) tavola sp. n. of the S. (R.) signata species group, but segment X has no spiny processes and the lateral lobes are situated very close mesally, not widely separated. Segment X is very similar to the unique, mesally angled lateral lobes at S. (R.) sarkoska sp. n. of the S. (R.) peruana species group.

87 88 89

Smicridea (Rhyacophylax) atmena sp. n., holotype, male genitalia: 85 = lateral view; 86 = left gonopod, ventral; 87 = dorsal; 88 = phallus lateral; 89 = phallus apex, ventral

Male genitalia. Segment IX subtriangular with high staked ventrum and produced anterolateral lobes in lateral view, anterodorsum indistinct; dorsal keel triangular tipped with microtrichiae; intersegmental profile between segments IX and X acute angled. Seg- ment X very long, lateral lobes almost tangential mesally producing long and very narrow mesal cleft; cerci longitudinally ovoid, located little nearer to basis; vestigial paraproct forming heavily sclerotised pair of internal straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopods, coxopodite high with very low basal fourth in lateral view; harpagones long slender parallel-sided slightly bending mesad and ventrad. Phallotheca with low basal portion

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)245 and arching stem; apex with asperites of ventromesal row of small spine points; a pair of dorsally serrated subapical ventrolateral clavate processes curving mesad; internal sclerites of sclerotized apical section of ejaculatory duct bipartite in ventral view.

Material examined – Holotype, male: : Chontachaca, Kosnipata-Cusco, 13°01’25” S, 71°28’03”W, 700 m, humid subtropical forest, Malaise trap, 13–30.VII.2005, leg. J. FAVIOLA MONTES CARLOS (1 male, NRM). Paratype: same data as holotype (1 male, OPC).

Etymology – Atmena, from “átmenõ”, transient in Hungarian, refers to mesally angled tips of the lateral lobes on segment X.

sp. n. (Figs 90–94)

Diagnosis – This species is close to S. (R.) fogasa sp. n. from Ecuador, from which it differs by having the ventrum IX without apicomesal process; the antecostal suture and paraproctal strap are separated in the strengthening frame; the dorsoapical process on the phallic apex encircling.

Description – Male. Pale brown, forewing length 6 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth darker and zigzag bordered; nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark.

93 92

Smicridea (Rhyacophylax) begorba sp. n., holotype, male genitalia: 90 = lateral view; 91 = left gonopod, ventral; 92 = dorsal; 93 = phallus lateral; 94 = phallus apex, ventral

Annls hist.-nat. Mus. natn. hung. 104, 2012 246 J. Oláh & K. A. Johanson

Male genitalia. Segment IX subtriangular with stalked ventrum, anterodorsum indistinct; dorsal keel triangular covered with microtrichiae; intersegmental profile between segments IX and X right angled. Segment X long, lateral lobes with inward curving and narrowing tip, bearing tiny setae; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral internal straps directed downward from apical margin; vestigial paraproct functioning as sclerotised phallic guides encircling above phallic organ; paraproctal straps separated from antecostal suture of segment IX. Basal segment of gonopods, coxopodite straight and reaching to tip of segment X; harpagones slender. Phallotheca with inflated vertical basal portion and apex with encircling dorsoapical processes and strong spinelike phallotremal sclerites.

Material examined – Holotype, male: : Amasonia, Gareno Lodge, Amaso- nian Lowland, 13–14.VIII.2011, light, leg. J. OLÁH Jr. (1 male, OPC).

Etymology – Begorba, from “begörbe”, inward curving in Hungarian, refers to the encircling dorsoapical process on the phallic apex.

Smicridea (Smicridea) bidentata MARTYNOV, 1912 (Figs 95–99)

Material examined – : Manu-Mother of Good, River Tonchaca, 71°20’W, 12° 50’S, 436 m, 17–22.V.2005, Malaise and light trap, leg. FAVIOLA MONTES CARLOS &JUAN CHAVEZ (1 male, NRM, 1 male, OPC). Chontachaca, Kosnipata-Cusco, 71°28’03’’ W, 13°01’25’’S, 700 m, Malaise trap 15–30.VII.2005, humid subtropical forest, leg. FAVIOLA MONTES CARLOS (19 males, 4 females, NRM, 19 males, 4 females, OPC). San Martin Prov., La Catarata de Ahuashiyascu, 6°27.544’S, 76°18.192’W, light, loc. 07, 7.I. 2009 leg. T. MALM & K. A. JOHANSON (1 male, NRM).

Remarks – Specimens collected in various habitats in Peru clearly exhibit the genital structures described and drawn very precisely by MARTYNOV (1912). In spite of the fact that most of his species drawings were prepared from uncleared terminalia, the visible detailes of the uncleared genital structures on his drawings are very indicative and reliable. Here we have redrawn the structure of the cleared genitalia.

Smicridea (Smicridea) bivittata (HAGEN, 1861)

Material examined – : Archidona, 1100 m, 9.VIII.2011,light, leg. J. OLÁH Jr. (1 male, OPC). Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 24.XI.2007, light, leg. J. OLÁH Jr. (2 males, OPC). Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 19.VIII.2011, light, leg. J. OLÁH Jr. (1 male, OPC). : State of Veracruz, Fortin de las Flores, Metlac River, under bridge of R 150, 900 mao, 24.VI.2006, light trap, leg. M. ESPELAND & T. MALM (3 males, NRM,

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)247

2 males, OPC). : San Martin Prov., Rio Negro, 37 km (rd.) W Moyobamba, near Olmos-Tarapoto rd., 6°00.278’S, 77°15.437’W, light, loc. 05, 6.I.2009, leg. K. A. JOHANSON & T. MALM (2 males, 2 females, NRM, 2 males, 1 female, OPC).

Remarks – S.(S.) bivittata has only a single subapical membrane band visible on specimens in alcohol, the subbasal band visible on pinned specimens has setal origin, not visible in setaless membrane of the specimens in alcohol.

95 96 97

98 99

Smicridea (Smicridea) bidentata MARTYNOV, 1912, male genitalia: 95 = lateral view; 96 = left gonopod, ventral; 97 = dorsal; 98 = phallus lateral; 99 = phallus ventral

Smicridea (Smicridea) calopa FLINT, 1974

Material examined – : State of Veracruz, Los Tuxtlas, around the Estacion the Biologia Los Tuxtlas, N 18°35.213’, W 095°04.462’, 30 mao, 27.VI.2006, net & hand pick, leg. M. ESPELAND & T. MALM (12 males, NRM, 8 males, OPC).

Smicridea (Smicridea) catherinae BLAHNIK, 1995

Material examined – : State of Veracruz, Fortin de las Flores, Metlac River, under bridge of R 150, 900 mao, 24.VI.2006, light trap, leg. M. ESPELAND &T.MALM (1 male, OPC). State of Veracruz, Los Tuxtlas area, Rio la Palma, near to the Estacion the Biologia Los Tuxtlas, N 18°33.680’, W 095°02.943’, 30 mao, 26.VI.2006, light trap, leg. M. ESPELAND &T.MALM (18 males, NRM, 12 males, OPC). State of Veracruz, Los Tuxtlas, around the Estacion the Biologia Los Tuxtlas, N 18°35.213’, W 095°04.462’, 30 mao, 27.VI.2006, net & hand pick, leg. M. ESPELAND &T.MALM (2 males, OPC). State of

Annls hist.-nat. Mus. natn. hung. 104, 2012 248 J. Oláh & K. A. Johanson

Veracruz, Los Tuxtlas area, Rio Maquinas, near to the Estacion the Biologia Los Tuxtlas, N 18°37.392’, W 095°05.418’, 9 mao, 27.VI.2006, light trap, leg. M. ESPELAND &T.MALM (7 males, NRM).

Smicridea (Rhyacophylax) columbiana (ULMER, 1905)

Material examined – : Misiones, PN, Iguazu, Research Station CIES, 185 m, 54°26.972’W, 25°40.733’S, 6.I.2008, leg. K. HILL,D.MARSHALL &M.MOULDS (1 male, NRM).

Smicridea (Rhyacophylax) discalis FLINT, 1972

Material examined – : Misiones, PN, Iguazu, Research Station CIES, 185 m, 54°26.972’W, 25°40.733’S, 6.I.2008, leg. K. HILL,D.MARSHALL &M.MOULDS (1 male, NRM).

Smicridea (Rhyacophylax) dithyra FLINT, 1974

Material examined – : State of Veracruz, Los Manantioles, Tlilapan, N 18°47.944’, W 097°06.270’, 1171 mao, 25.VI.2006,light trap, leg. M. ESPELAND &T.MALM (10 males, 150 females, NRM, 4 males, 10 females, OPC). State of Veracruz, Fortin de las Flores, Metlac River, under bridge of R 150, 900 mao, 24.VI.2006, light trap, leg. M. ESPELAND & T. MALM (1 male, NRM).

sp. n. (Figs 100–104)

Diagnosis – This species belongs to the S.(S.) fasciatella species group, and is close to S.(S.) erecta FLINT, 1974 from Surinam, from which it is distinguished by the less produced and less erect angle of the apicomesal lobe; small cerci and depigmented window pattern different; the harpagones very thin; the stem of the phallotheca slightly arching convex dorsad, not straight or concave in lateral view.

Description – Male. Dark brown, forewing length 4 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands without long anterolateral filament, but with 2 pairs of internal reticulate sacs in subapical abdominal segments. Forewing membrane dark homogenous with a faint indication of a pale band at stigmal region.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)249

101

100

103 102 104

Smicridea (Smicridea) dombora sp. n., holotype, male genitalia: 100 = lateral view; 101 = left gonopod, ventral; 102 = dorsal; 103 = phallus lateral; 104 = phallus apex, ventral

Male genitalia. Segment IX with low and long basodorsal excision. Segment X enlarged, lateral processes rounded; apicomesal corner produced upward; cerci small located dorsad of depigmented window; vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin, less developed. Basal segment of gonopods, coxopodite stout; harpagones thin. Phallotheca with dorsad arching stem.

Material examined – Holotype, male: : Approuaguekaw, Kaw Mt, 104 mao, 4°33.035’N, 52°11.661’W, Malaise trap, 23.I–7.II.2007, FRG MF3, leg. N. JÖNSSON (NRM). Paratypes: same data as holotype (2 males, NRM). Approuaguekaw, Kaw Mt, 104 m, 4°33.035’N, 52°11.661’W, Malaise trap, 4–12.II.2007, FRG MF3, leg. N. JÖNSSON (1 male, NRM, 1 male, OPC).

Etymology – Dombora, coined from “domború”, convex in Hungarian, refers to the dorsad arching, convex stem of the phallotheca.

sp. n. (Figs 105–109)

Diagnosis – This species is a member of the S. (S.) nigripennis group, and related to S. (S.) rara BUENO-SORIA et MÁRQUEZ-MAJAUDON, 1979 from Mexico, from which it is distinguished by having no lateral subapical spines, but having basolateral spines on the ventrolateral processes of the phallotheca.

Annls hist.-nat. Mus. natn. hung. 104, 2012 250 J. Oláh & K. A. Johanson

Description – Male. Dark, almost black forewing length 4 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands without long anterolateral filament, but with 2 pairs of internal reticulate sacs in subapical abdominal segments. Forewing membrane dark homogenous with lighter area on pterostigma and arculus. Male genitalia. Segment IX triangularly produced anterad; intersegmental profile between segments IX and X almost indistinct. Segment X short, lateral lobes truncate in dorsal view; cerci small represented by a few setae; vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin. Basal segment of gonopods, coxopodite expanded slightly apicad; harpagones long slender parallel-sided slightly pointed in lateral view. Phallotheca with ventroapiocal processes having laterad curving tips and without lateral subapical spine.

Material examined – Holotype, male: : State of Veracruz, Tecomaluca, Sierra de agua, N 18°44.977’, W 097°14.576’, 1420 mao, 25.VI.2006, net and hand pick, leg. M. ESPELAND &T.MALM (NRM). Paratypes: same data as holotype (1 male, NRM, 1 male, OPC).

Etymology – Egsera, coined from “egyszerû”, simple in Hungarian, refers to the simple apical portion of the ventroapical processes of phallotheca, without robust laeral subapical spine.

105 106

107 108

109

Smicridea (Smicridea) egsera sp. n., holotype, male genitalia: 105 = lateral view; 106 = left gonopod, ventral; 107 = dorsal; 108 = phallus lateral; 109 = phallus ventral

Smicridea (Rhyacophylax) ephippifera FLINT, 1978

Material examined – : St-Laurentdu-Maroni: Maroni River: A- Chimato-Campop, village 5°13.716’N, 54°17.498’W, 6.II.2007, FRG 12, leg. N. JÖNSSON (1 male, NRM).

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)251

sp. n. (Figs 110–114)

Diagnosis – This species is a member of the S. (S.) nigripennis species group, and is most similar to S.(S.) appendicula FLINT, 1974 from Surinam, from which it is distinguished by having segment IX high, not low; the anterodorsum of segment IX is excised, not straight; the segment X is short, not long; the harpagones are slender, not stout; the phallotheca is differently shaped; the phallotheca is covered by short spines, not bare.

Description – Male. Dark brown, forewing length 4 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands without long anterolateral filament, but with 2 pairs of internal reticulate sacs in subapical abdominal segments. Forewing membrane dark homogenous without any pattern. Male genitalia. Segment IX with basodorsal excision. Segment X short, lateral processes blunt; cerci elongated; vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin, less developed. Basal segment of gonopods, coxopodite stout; harpagones thin. Phallotheca with a single dorsal median process and deeply cleft ventrum; endotheca with a pair of S-formed and a pair of shorter straight spines larger apical half of phallotheca covered with short spines.

Material examined – Holotype, male: : Approuaguekaw, Kaw Mt, 104 mao, 4°33.035’N, 52°11.661’W, Malaise trap, 23.I–7.II.2007, FRG MF3, leg. N. JÖNSSON (NRM). Paratypes: same data as holotype (6 males, NRM, 2 males, OPC). Approuague- kaw, Kaw Mt, 104 m, 4°33.035’N, 52°11.661’W, Malaise trap, 4–12.II.2007, FRG MF3, leg. N. JÖNSSON (7 males, NRM, 4 males, OPC).

Etymology – Erda, coined from “érdes”, rough in Hungarian, refers to the short spine- covered surface of the phallotheca.

110

114

112

113 111 Smicridea (Smicridea) erda sp. n., holotype, male genitalia: 110 = lateral view; 111 = left gonopod, ventral; 112 = dorsal; 113 = phallus ventral; 114 = phallus lateral

Annls hist.-nat. Mus. natn. hung. 104, 2012 252 J. Oláh & K. A. Johanson

Smicridea (Smicridea) erecta FLINT, 1974

Material examined – : Approuaguekaw, Kaw Mt, 216 m, 4°33.257’ N, 52°11.920’W, Malaise trap, 19.I–4.II.2007, FRG MF 2, leg. N. JÖNSSON (3 males, OPC). Approuaguekaw, Kaw Mt, 104 mao, 4°33.035’N, 52°11.661’W, Malaise trap, 23.I–7.II. 2007, FRG MF3, leg. N. JÖNSSON (1 male, NRM).

sp. n. (Figs 115–119)

Diagnosis – This species having apicodorsal phallic lobes belongs to the S.(R.) murina species group, and is closest to S.(R.) murina MCLACHLAN, 1871, but differs by having lateral lobes on segment X mesad directed, not posterad; gonopods parallel-sided almost along entire length, not constricted basad; dorsoapical processes on phallic apex long and straight, not hornlike.

116

115

117 118 119

Smicridea (Rhyacophylax) felsa sp. n., holotype, male genitalia: 115 = lateral view; 116 = left gonopod, ventral; 117 = dorsal; 118 = phallus lateral; 119 = phallus apex, ventral

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)253 angled. Lateral lobes on segment X with mesad directed tip; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral straps directed downward from apical margin and fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite straight, parallel-sided and reaching to tip of segment X; harpagones slender parallel-sided slightly bending mesad. Phallotheca with low vertical inflated basal portion; apex with a pair of distinct dorsoapical long processes and sclerotised apical half of ejaculatory duct, ending in phallotremal sclerites like a pair of spines.

Material examined – Holotype, male: : Amasonian Lowland, Gareno, near Puerto Napo, 400 m, 14.VIII.2011, light, leg. J. OLÁH Jr. (OPC).

Etymology – Felsa, from “felsõ”, upper in Hungarian, refers to the dorsoapical phallic processes.

sp. n. (Figs 120–124)

Diagnosis – This species having spiny process on the middle of the ventrolateral margin of the paraproct forms a closely related species complex together with S. (R.) salta FLINT, 1974, S.(R.) nemorosa HOLZENTHAL et BLAHNIK, 1995 and S.(R.) kampoka sp. n. Closest to S. (R.) nemorosa from Costa Rica, but easily distinguished by the presence of short ventromesal posterior triangular lobe on stermun IX, absent at S. nemorosa; the clearly inward curving and narrowing tip of the lateral lobes on segment X, not with blunt apex; coxopodite angulate ventrobasad, not straight; harpagones capitate only in lateral view, narrowing in ventral view; apex of phallic organ differently structured.

Description – Male. Pale brown, forewing length 4.5 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth light; nygmal and thyridial points as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. Male genitalia. Segment IX subtriangular with stalked ventrum, anterodorsum indistinct; dorsal keel rounded triangular covered with microtrichiae; intersegmental profile between segments IX and X sharp rounded angled; ventromesally with a posterior short triangular lobe. Segment X long, lateral lobes with inward curving and narrowing tip, bearing tiny setae; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral internal straps directed downward from apical margin; ventrolateral margin of segment X bearing single spinelike projection; vestigial paraproct functioning as sclerotised phallic guides encircling above phallic organ; paraproctal straps fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite straight and reaching to tip of segment X; harpagones slender capitate in lateral and narrowing in ventral view. Phallotheca with inflated vertical basal portion and apex with plumose headed lateral processes and 2 pairs of internal spines.

Annls hist.-nat. Mus. natn. hung. 104, 2012 254 J. Oláh & K. A. Johanson

Material examined – Holotype, male: Wild Sumaco, near Pacto Sumaco, 7–8.X.2010, light, leg. J. OLÁH Jr. (OPC).

Etymology – Fogasa, from “fogas”, dentate in Hungarian, refers to the dentate, almost plumose headed lateral processes on the phallic apex.

121

120

122 123

124

Smicridea (Rhyacophylax) fogasa sp. n., holotype, male genitalia: 120 = lateral view; 121 = left gonopod, ventral; 122 = dorsal; 123 = phallus lateral; 124 = phallus apex, ventral

Smicridea (Smicridea) frequens (NAVAS, 1930)

Material examined – : Region del Biobio (VIII), ca. 20 km WSW, Arauco, stream crossing road at Puento Caripilum, 37°18.141’S, 73°30.633’W, 20 m (Loc #06), light trap 2.I.2006, K. A. JOHANSON (16 males, NRM, 4 males, OPC). Region de los Lagos (X), PN, Vicente Rosales, 200 m downstream Saltos de Petrohué, 6 km SW, Petrohué and Lago Todo los Santos, stream parallel to Rio Petrohué, 41°10.378’S, 72°27.057’W, 110 m (Loc #18), light trap 12.I.2006, K. A. JOHANSON (2 males, NRM, 3 males, OPC). Region del Maule (VII), river ca. 43.4 km ESE Curicó, Potrero Grande, at Puente los Morongos, 35° 12.990’ S, 70°58.517’W, 727 m (Loc #31), light trap 20.I.2006, K. A. JOHANSON (5 males, NRM).

sp. n. (Figs 125–129)

Diagnosis – This species having apicodorsal phallic lobes belongs to S.(R.) murina species group, and closest to S. (R.) murina MCLACHLAN, 1871, from which it differs by

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)255 having more developed sutures on segment IX; ventral stem of segment IX very short; dorsoapical horn on phallic apex thick; a pair of serrulated lines present on the ventrum of phallic apex.

Description – Male. Pale brown, forewing length 4.8 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: white transverse band at apical fourth; chord (anastomosis) of crossveins s, r-m, m, m-cu and cu darker. Male genitalia. Segment IX subtriangular with stalked ventrum, strap of stalk is reduced to antecostal suture fused with paraproct; anterodorsum indistinct; dorsal keel rounded; intersegmental profile between segments IX and X obtuse angled. Segment X short, lateral lobes with rounded tip; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral straps directed downward from apical margin and fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite straight and reaching to tip of segment X; harpagones slender parallel-sided slightly bending mesad. Phallotheca with vertical inflated basal portion almost as high as length of stem; apex with distinct dorsoapical thick horns and sclerotised apical half of ejaculatory duct, ending in phallotremal sclerites like a pair of spines, a pair of serrulated lines present subapicad on ventrum of phallic apex.

120

125

128 127

129

Smicridea (Rhyacophylax) furesa sp. n., holotype, male genitalia: 125 = lateral view; 126 = left gonopod, ventral; 127 = dorsal; 128 = phallus lateral; 129 = phallus apex, ventral

Material examined – Holotype, male: : Archidona, 1100 m, 09.VIII.2011, light, leg. J. OLÁH Jr. (1 male, OPC). Paratypes: same data as holotype (1 male, 1 female, OPC). : Manu-Mother of Good, River Yunguya 71°22’W, 12°49’S, 546 m, 9–15.V.2005,

Annls hist.-nat. Mus. natn. hung. 104, 2012 256 J. Oláh & K. A. Johanson

Malaise and light trap, leg. FAVIOLA MONTES CARLOS &JUAN CHAVEZ (50 males, 50 females, NRM, 30 males, OPC). Manu-Mother of Good, River Gamitana 71°21’W, 12°52’S, 546 m, 24–30.V.2005, Malaise and light trap, leg. FAVIOLA MONTES CARLOS & JUAN CHAVEZ (100 males, 100 females, NRM). Palotoa, Manu-Mother of Good, 71°18’W, 12°35’S, 436 m, 24–30.V.2005, Malaise and light trap, leg. FAVIOLA MONTES CARLOS & JUAN CHAVEZ (50 males, 50 females, NRM). San Martin Prov., Rio Huallaga, at Pumarihri Huallaga Lodge, between Juan Guerra and Chazuta, 14 km (rd) W Chazuta, 6°36.643’S, 76°12.555’W, light, loc. 10, 9.I.2009, leg. K. A. JOHANSON &T.MALM (1 male NRM). San Martin Prov., Rio Huallaga tributary, small river passing Chazuta, 6°34.665’S, 76°08.209’ W, light, loc. 11, 10.I.2009, leg. T. MALM & K. A. JOHANSON (1 male, NRM).

Etymology – Furesa, from “fûrészes”, serrated in Hungarian, refers to the pair of serrulate lines developed subapicad on the phallic apex.

sp. n. (Figs 130–134)

Diagnosis – This species belongs to the S. (R.) signata group, and is close to S. (R.) pseudolobata FLINT, 1978 from Brazil, from which it is distinguished in dorsal view by the mesad curving tip of the lateral lobes of segment X, not straight both in lateral and dorsal view; coxopodite stalked, not parallel-sided; apex of phallic organ differently shaped.

Description – Male. Pale brown, forewing length 4.3 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: white transverse band at apical fourth; chord (anastomosis) of crossveins s, r-m, m, m-cu and cu darker. Male genitalia. Segment IX subtriangular with vertical stalk, anterodorsum indistinct; dorsal keel rounded; intersegmental profile between segments IX and X indistinct. Segment X short, lateral lobes with mesad curving tip; a pair of serrated and spiny apicoventral lobes present; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of internal straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ; paraproctal straps confused with antecostal suture of segment IX forming together an Y-shaped strengthening frame, a rather well-sclerotised strap forming a dorsal guide on phallic organ. Basal segment of gonopods, coxopodite stalked and straight and reaching to tip of segment X; harpagones slender parallel-sided slightly bending mesad and ventrad. Phallotheca with low vertical inflated basal portion and apex with serrated and membranous lateral lobes; ventral keel accompanied by short serrated ridges.

Material examined – Holotype, male: : Amasonian Lowland, Gareno, near Puerto Napo, 10–11.X.2010, light, leg. J. OLÁH Jr. (OPC).

Etymology – Hajla from “hajló” curving in Hungarian, refers to the mesad curving tips of the lateral lobes on segment X.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)257

131

130

132 134

133

Smicridea (Rhyacophylax) hajla sp. n., holotype, male genitalia: 130 = lateral view; 131 = left gonopod, ventral; 132 = dorsal; 133 = phallus lateral; 134 = phallus apex, ventral

sp. n. (Figs 135–139)

Diagnosis – This species has branched apex on the harpagones as in S. bivittata (HAGEN, 1861) and S. bidentata MARTYNOV, 1912. The more robust and short coxopodite, as well as the more complex segment X relates this species to members in the species complex with S. breviuncata FLINT, 1974, S. conjuncta FLINT, 1991, S. nigerrima FLINT, 1983, S. prorigera FLINT, 1991, S. ruginasa FLINT, 1991, S. saucia MCLACHLAN, 1871 and S. urra FLINT, 1991. It is most similar to S. (S.) sarla sp. n. from Peru, from which it differs by having harpagones with 3-branched apex, not simple tapering; phallic organ and segment X differently shaped.

Description – Male. Almost uniformly fuscous with indistinct and disjunct transversal band little basad of forewing middle; forewing length 5 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands with long anterolateral filament, and 2 pairs of internal reticulate sacs present in subapical abdominal segments. Male genitalia. Segment IX produced convex on middle anterad; intersegmental profile between segments IX and X right angled. Segment X with rather unusual bilobed specific shape; cerci present on mesal lobe; vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin. Basal segment of gonopods, coxopodite robust and short, harpago three-branched. Phallotheca with bilobed apex.

Annls hist.-nat. Mus. natn. hung. 104, 2012 258 J. Oláh & K. A. Johanson

Material examined – Holotype, male: : San Martin Prov., La Catarata de Ahua- shiyascu, 6°27.544’S, 76°18.192’W, light, loc. 07, 7.I.2009, leg. T. MALM &K.A.JOHANSON, (NRM).

Etymology – Haraga, coined from “háromágas”, three-branched in Hungarian, refers to the three-branched apex of the harpagones.

135 136

139

137 138

Smicridea (Smicridea) haraga sp. n., holotype, male genitalia: 135 = lateral view; 136 = left gonopod, ventral; 137 = dorsal; 138 = phallus lateral; 139 = phallus apex, ventral

sp. n. (Figs 140–144)

Diagnosis – With the three dark points dorsoapically on the phallic apex, the species belongs to S. (R.) argentina species group. The slender phallotheca relates it to the S.(R.) argentina species complex. The three dark points located almost on the same coronal plane, not in different level like at S. argentina (NAVÁS, 1918) and S. palmar SGANGA, 2005. More- over in dorsal view the lateral margin of the lateral lobes on segment X concave, not convex resulting in a pair of mesal digitate processes variously adhered to each others on cleared animals.

Description – Male. Pale brown, forewing length 67 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth darker and zigzag bordered; nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)259

Male genitalia. Segment IX subtriangular with high staked ventrum and produced anterolateral lobes in lateral view, anterodorsum indistinct; dorsal keel rounded covered with microtrichiae; intersegmental profile between segments IX and X right angled. Seg- ment X very long, lateral lobes with concave lateral margin in dorsal view; cerci longitudinally ovoid; vestigial paraproct forming heavily sclerotised pair of internal straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopods, coxopodite high with very low basal fourth in lateral view; harpagones long slender parallel-sided slightly bending mesad and ventrad with pointed apex in ventral view. Phallotheca with high basal portion and straight stem; apex with dorsal subapical hump in lateral view; 3 dark points located on similar level of coronal plane; erectible endotheca with a pair of short dark hooks and erectible endophallus with sclerous band of internal sclerites having dilated apical section.

Material examined – Holotype, male: : Amasonian Lowland, Gareno, near Puerto Napo, 400 m, 14.VIII.2011,light, leg. J. OLÁH Jr. (OPC). Paratypes: : San Mar- tin Prov., Rio Huallaga tributary, small river passing Chazuta, 6°34.665’S, 76°08.209’ W, light, loc. 11, 10.I.2009, leg. T. MALM & K. A. JOHANSON (2 males, NRM, 1 male, OPC).

Etymology – Homora, from “homorú”, concave in Hungarian, refers to the lateral lobes of segment X with concave apicolateral margin in dorsal view.

sp. n. (Figs 145–149)

Diagnosis – This new species is a member of the S.(S.) nigripennis group, and differs from all known species by having unique double-hookedlateral processes on the phallotheca.

Description – Male. Dark, almost black forewing length 5 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands without long anterolateral filament, but with 2 pairs of internal reticulate sacs of pheromonal glands in subapical abdominal segments. Forewing membrane dark homogeneous with lighter transversal band a little anterad of chord. Male genitalia. Segment IX produced anterad; intersegmental profile between segments IX and X almost indistinct. Segment X short, lateral lobes rounded; cerci small represented by a few setae; vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin. Basal segment of gonopods, coxopodite slender, expanded slightly apicad; harpagones long slender parallel-sided clavate in lateral view. Phallotheca with closely appressed straight apical and double-hooked lateral processes.

Material examined – Holotype, male: : Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 24.XI.2007, light, leg. J. OLÁH Jr. (OPC).

Etymology – Horga, from “horgas”, barbed in Hungarian, refers to the double-hooked lateral processes on the phallotheca.

Annls hist.-nat. Mus. natn. hung. 104, 2012 260 J. Oláh & K. A. Johanson

141 140

143

142

144

Smicridea (Rhyacophylax) homora sp. n., holotype, male genitalia: 140 = lateral view; 141 = left gonopod, ventral; 142 = dorsal; 143 = phallus lateral; 144 = phallus lateral, erected (paratype)

145 146

147

148 149

Smicridea (Smicridea) horga sp. n., holotype, male genitalia: 145 = lateral view; 146 = left gonopod, ventral; 147 = dorsal; 148 = phallus lateral; 149 = phallus ventral

Smicridea (Smicridea) inaequispina FLINT, 1974

Material examined – : Approuaguekaw, Kaw Mt, 4°33.266’N, 52° 11.898’W, 249 mao, 17.I.2007, FRG 1, leg. N. JÖNSSON (3 females, NRM, 1 male, 1 female; OPC).

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)261

sp. n. (Figs 150–154)

Diagnosis – A member of S. (R.) salta species complex. Closest to S. (R.) fogasa sp. n., but easily distinguished by the absence of short ventromesal posterior triangular lobe; S. (R.) nemorosa lacking this lobe, but the ventrolateral and ventromesal lobes as well as the slender sclerotised rods on the phallic apex differently formed.

Description – Male. Pale brown, forewing length 4 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: subapical transversal band light; nygmal and thyridial points as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. Male genitalia. Segment IX subtriangular with stalked ventrum, anterodorsum indistinct; dorsal keel rounded triangular covered with microtrichiae; intersegmental profile between segments IX and X right rounded angled; ventromesally without posterior triangular lobe. Segment X long, lateral lobes with inward curving and narrowing tip, bearing tiny setae; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral internal straps directed downward from apical margin; ventrolateral margin of segment X bearing single spinelike projection; paraproctal straps fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite straight and reaching to tip of segment X; harpagones slender slightly capitate. Phallotheca with inflated vertical basal portion and apex with fingered ventrolateral and blunt ventromesal lobes; internal recractile structure composed of a pair of long spinelike rods hooked laterad and sclerotised ejaculatory duct with indistinct hook inside. 151

150 154 152 153

Smicridea (Rhyacophylax) kampoka sp. n., holotype, male genitalia: 150 = lateral view; 151 = left gonopod, ventral; 152 = dorsal; 153 = phallus lateral; 154 = phallus apex, ventral

Annls hist.-nat. Mus. natn. hung. 104, 2012 262 J. Oláh & K. A. Johanson

Material examined – Holotype, male: : San Martin Prov., creek crossing rd. Tarapoto-Yurimaguas, ca. 30 km (rd.) NE Tarapoto, 6°24.904’S, 76°18.756’W, light, loc. 08, 8.I.2009, leg. K. A. JOHANSON &T.MALM (NRM). Paratypes: same data as holotype (2 males, NRM, 1 male, OPC).

Etymology – Kampoka, from “kampóka”, diminutive hook in Hungarian, refers to the pair of long spinelike hooked rods on the phallic apex.

sp. n. (Figs 155–159)

Diagnosis – This species belongs to the S.(S.) fasciatella species group, and is close to S.(S.) aequalis BANKS, 1920 from Guiana, but easily distinguished by having harpagones spatulate, not simple tapering.

Description – Male. Dark, almost black without any membrane pattern; forewing length 5 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands without long anterolateral filament, but with 2 pairs of internal reticulate sacs in subapical abdominal segments.

156

155 158 157

159

Smicridea (Smicridea) kana sp. n., holotype, male genitalia: 155 = lateral view; 156 = left gonopod, ventral; 157 = dorsal; 158 = phallus lateral; 159 = phallus apex, ventral

Male genitalia. Segment IX convex anterad; intersegmental profile between segments IX and X almost absent. Segment X enlarged, lateral processes slightly upward produced apically in lateral view and tapering in dorsal view; cerci large located laterally subapicad;

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)263 vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin, its basal part more developed. Basal segment of gonopods, coxopodite high, low in basal third; harpagones spatulate. Phallotheca with low basal section and with almost parallel-sided stem and rounded membranous apex.

Etymology – Kana, coined from “kanál”, spoon in Hungarian, refers to the spoon- shaped apex of the harpagones.

sp. n. (Figs 160–164)

Diagnosis – Smicridea (R.) radula FLINT, 1991, S. (R.) pseudoradula FLINT, 1991 and S. (R.) kapara sp. n. are very simiar. Having short and blunt lateral lobes on segment X accompanied ventrolaterally by the heavily sclerotised paraproct with apex of pointed projection they are forming a species complex agreeing in most structures, except the phallic organ. Dorsal lobe of the phallic apex short and narrow quadratic at S. pseudoradula, long and broad quadratic at S. radula, but long and rounded with constricted basal stalk at S. kapara. Moreover the new species has no any sign of basolateral spiculate patch present at S. radula and S. pseudoradula.

160 161

163

164 162

Smicridea (Rhyacophylax) kapara sp. n., holotype, male genitalia: 160 = lateral view; 161 = left gonopod, ventral; 162 = dorsal; 163 = phallus lateral; 164 = phallus apex, ventral

Annls hist.-nat. Mus. natn. hung. 104, 2012 264 J. Oláh & K. A. Johanson

Description – Male. Small pale brown animal, forewing length 4 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane without distinct pattern, except crossveins m-cu and cu more pronounced and pigmented. Male genitalia. Segment IX with anterolateral margin produced as a narrow, slightly upturned lobe; sternum with a narrow, pointed process between gonopod bases; intersegmental profile between segments IX and X acute angled. Segment X short, lateral lobes short and blunt; cerci rounded ovoid, located apicad. Vestigial paraproct forming heavily sclerotised pair of ventrolateral internal straps with apical apex projecting with several small points; vestigial paraproct functioning as sclerotised phallic guides encircling above phallic organ; paraproctal straps fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite straight and reaching over tip of segment X; harpagones slender. Phallotheca with inflated vertical basal portion and apex with rounded dorsal stalked lobe and a pair of ventrolateral lobes armed with a series of short, dark teeth recurving basad.

Material examined – Holotype, male: : Chontachaca, Kosnipata-Cusco, 13° 01’25”S, 71°28’03”W, 700 m, humid subtropical forest, Malaise trap 13–30.VII.2005, leg J. FAVIOLA MONTES CARLOS (NRM). Paratypes: same data as holotype (3 males NRM, 3 males OPC). San Martin Prov., creek crossing rd. Juan Guerra-Chazuta, 14 km (rd.) E Colombia Bridge, 6°35.594’S, 76°13.172’W, light, loc. 09, 9.I.2009, leg. K. A. JOHANSON & T.MALM (1 male, NRM).

Etymology – Kapara, from “kaparó”, scraper in Hungarian, refers to the dentate radula-like ventrolateral lobes on the phallic apex.

sp. n. (Figs 165–169)

Diagnosis – This species belongs to the S.(S.) fasciatella species group, and is similar to S.(S.) bidentata MARTYNOV, 1912 from Peru, but easily distinguished by having anterior margin of segment IX produced triangularly, not simple convex; bifid apex of harpago less excised; phallotheca broad “fat” in lateral view, not slender apex of phallotheca broadening in ventral view, not capitate.

Description – Male. Dark brown, forewing length 5 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands without long anterolateral filament, but with 2 pairs of internal reticulate sacs in subapical abdominal segments. Forewing membrane dark homogenous with faint hyaline window along stem M and at arculus. Male genitalia. Segment IX with triangularly produced anterior margin. Segment X elongated, lateral processes pointed; apicomesal corner produced slightly upward; cerci small located dorsad of depigmented window; vestigial paraproct forming sclerotised pair

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)265 of lateroventral straps directed downward from apical margin, less developed. Basal segment of gonopods, coxopodite stout; harpagones shallow bifid. Phallotheca broad, slighly arching upward.

Material examined – Holotype, male: : Jujuy, RN, 9, nr. Leon, 1693 m, 24°01.9565’S, 65°26.519’W, 17.I.2008, leg. K. HILL,D.MARSHALL &M.MOULDS (1 male, NRM).

Etymology – Kovera, coined from “kövér”, fat in Hungarian, refers to the broad phallotheca. 165 166

168

167 169

Smicridea (Smicridea) kovera sp. n., holotype, male genitalia: 165 = lateral view; 166 = left gonopod, ventral; 167 = dorsal; 168 = phallus lateral; 169 = phallus apex, ventral

sp. n. (Figs 170–174)

Diagnosis – This species belongs to S.(R.) peruana species group, and is close to S. (R.) andicola FLINT, 1991 from Colombia, but differs by having long pointed microtrichous dorsal keel on tergum IX, not short and blunt; apicomesal process of sternum IX long, not short; lateral lobes on segment X turning mesad, not posterad directed; their apical points short, not long; apicolateral surfaces of the phallic apex with elongated asperite patch of spicules, absent at andicola.

Annls hist.-nat. Mus. natn. hung. 104, 2012 266 J. Oláh & K. A. Johanson nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. Male genitalia. Segment IX subtriangular, anterodorsum indistinct; dorsal keel long triangular tipped with microtrichiae; intersegmental profile between segments IX and X acute angled with a small membraneous process below tip of dorsal keel; apicoventral lobe on sternum IX long. Segment X elongated, lateral lobes curving mesad with minute pointed apex; its ventrolateral margin with a small sclerotised lobe; cerci longitudinally ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of internal straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopods, coxopodite high with low basal fourth in lateral view; harpagones long slender, parallel-sided and slightly bending mesad and ventrad. Phal- lotheca with vertical inflated basal portion right angled to horizontal stem; phallic apex with lateral elongated patches and a single apicoventral row of small spine points; a pair of ventrolateral serrated lobes present apicad; ejaculatory duct indistinct.

Material examined – Holotype, male: : Amazonas Prov., river crossing Olmos- Tarapoto rd., 371 km (rd.) E Olmos Desv. Jaén, 5°41.178’S, 77°46.421’W, light, loc. 02,. Paratypes: same data as holotyp 5.I.2009, leg. K. A. JOHANSON &T.MALM (NRM)e (10 males, NRM, 6 males, OPC). San Martin Prov., river crossing rd. Olmos-Tarapoto, 385 km (rd.) E Olmos Desv. Jaén, 5°40.055’S, 77°43.396’W, light, loc. 04, 5.I.2009, leg. K. A. JOHANSON &T.MALM (1 male, OPC).

Etymology – Lebena, from “lebeny”, lobe in Hungarian, refers to the lobe developed on both sides of the ventrolateral margin of segment X.

170 171

173

172

174

Smicridea (Rhyacophylax) lebena sp. n., holotype, male genitalia: 170 = lateral view; 171 = left gonopod, ventral; 172 = dorsal; 173 = phallus lateral; 174 = phallus apex, ventral

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)267

sp. n. (Figs 175–178)

Diagnosis – By having a unique fanlike structure with spiny margin somehow connected to paraproct and functioning as phallic guide, this species is similar to S.(R.) ephippifera FLINT, 1978 and S.(R.) dentiserrata ALBINO,PES et HAMADA, 2011. This structure is however fanlike at S.(R.) legeza sp. n., not elongated lobelike; the lateral lobe on segment X is rounded, not pointed or truncate.

Description – Male. Pale brown, forewing length 5 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth dark with zigzag margin; chord (anastomosis) of crossveins s, r-m, m, m-cu and cu darker. Male genitalia. Segment IX subtriangular with stalked ventrum; anterodorsum indistinct; dorsal keel rounded triangular in dorsal view; intersegmental profile between segments IX and X flat. Lateral lobes on segment X with blunt in dorsal and upward angled in lateral view; cerci rounded ovoid, located middle; vestigial paraproct forming sclerotised pair of ventrolateral straps directed downward from apical margin and fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame; a fanlike structure with spiny margin connected to paraproct and forming a phallic guide. Basal segment of gonopods, coxopodite straight, parallel-sided and slightly surpassing tip of segment X; harpagones slender parallel-sided slightly bending mesad. Phallotheca with low vertical basal portion; apex with endotheca in retracted position exhibiting dubious lobes, long endophallus and short internal sclerite, structure even more indistinct in ventral view.

176

175

177 178

Smicridea (Rhyacophylax) legezoa sp. n., holotype, male genitalia: 175 = lateral view; 176 = left gonopod, ventral; 177 = dorsal; 178 = phallus lateral

Annls hist.-nat. Mus. natn. hung. 104, 2012 268 J. Oláh & K. A. Johanson

Material examined – Holotype, male: : San Martin Prov., Rio Negro, 37 km (rd.) W Moyobamba, near Olmos-Tarapoto rd., 6°00.278’S, 77°15.437’W, light, loc. 05, 6.I. 2009, leg. K. A. JOHANSON & T. MALM (NRM).

Etymology – Legezoa, from “legyezõ”, fan in Hungarian, refers to the unique fanlike structure of the paraproct.

sp. n. (Figs 179–183)

Diagnosis – This species, with spiny process on the ventrolateral margin of the paraproct, belongs to the S.(R.) signata species group, and is closest to S.(R.) bidactyla FLINT et REYES-ARRUNATEGUI, 1991, but easily distinguished by having paraproctal spine complex tridactylate, not bidactylate; apices of segment X with 3 points, not with 2 points; 2 patches of spines present dorsolaterad on the middle of phallic stem, not a single saddle-shaped; midventral row of spines absent; ventrolateral short spiny lobes and subquadratic structure bearing anterad directed short lateral tooth present on phallic apex.

Description – Male. Pale brown, forewing length 4.5 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth dark marked by transversal darker zigzag and lighter band; nygmal and thyridial points as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. Male genitalia. Segment IX subtriangular with stalked ventrum, anterodorsum indistinct; dorsal keel rounded covered with microtrichiae; intersegmental profile between segments IX and X acute angled. Segment X long, lateral lobes with inward curving pointed tip and mesal subapical spine; cerci rounded ovoid, located middle; paraproct forming heavily sclerotised pair of ventrolateral straps on ventrolateral margin of segment X bearing tridactylate spine complex subapicad; very pronounced long branch directed downward deep; 2 small spines directed dorsad and mesad; paraproct functioning as sclerotised phallic guides encircling above phallic organ; paraproctal straps fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite slightly S-forming and reaching to tip of segment X; harpagones slender. Phallotheca with inflated low vertical basal portion and horizontal stem with 2 spiny patches on midway dorsolaterad; apex with semierect dorsal bilobed plate and narrow capitate ventromesal tonguelike lobe accompanied by a pair of short lateral lobe with spiny apex; recractile internal sclerite bilobed; subquadratic structure visible somehow attached dorsoapicad bearing anterad a pair od short spines.

Material examined – Holotype, male: : San Martin Prov., Rio Mayo, 11 km (rd.) E Mayobamba, 6°04.989’S, 76°53.065’W, light, loc. 06, 6.I.2009, leg. T. MALM &K.A. JOHANSON (NRM). Paratypes: same data as holotype (5 males, NRM, 3 males, OPC).

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)269

180 179

182 181

183

Smicridea (Rhyacophylax) leloga sp. n., holotype, male genitalia: 179 = lateral view; 180 = left gonopod, ventral; 181 = dorsal; 182 = phallus lateral; 183 = phallus apex, ventral

Etymology – Leloga, from “lelógó”, hanging down in Hungarian, refers to the enlarged spine of paraproct directed downward on the ventrolateral margin of segment X.

sp. n. (Figs 184–188)

Diagnosis – This new species is a member of the S. (S.) nigripennis species group, and closest to S.(S.) inaequispina FLINT, 1974 from Surinam, but differs by having lateral pattern of the combined paraproct and suture brace complex completely different; indis- cernible any dorsomesal process on phallotheca; 6 endothecal spines with similar length.

Description – Male. Dark brown, forewing length 4 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands with halfsegment long anterolateral filament, 2 pairs of internal reticulate sacs in subapical abdominal segments present. Forewing membrane dark homogenous with hyaline window pattern only present along stem M1 and on Cu2 confluence at arculus. Male genitalia. Segment IX with long basodorsal excision in lateral view. Segment X short, lateral processes bilobed, mesal lobe smaller; cerci elongated setose surface; vestigial paraproct of sclerotised pair of lateroventral straps directed downward from apical margin. Basal segment of gonopods, coxopodite stout; harpagones long with narrowing apices. Phallotheca with a single long ventromesal processes; endotheca with 3 pairs of stout and long spines.

Annls hist.-nat. Mus. natn. hung. 104, 2012 270 J. Oláh & K. A. Johanson

184 185

187

188

186

Smicridea (Smicridea) lemeza sp. n., holotype, male genitalia: 184 = lateral view; 185 = left gonopod, ventral; 186 = dorsal; 187 = phallus lateral; 188 = phallus ventral

Material examined – Holotype, male: : San Martin Prov., stream crossing Juan Guerra-Chazuta rd., 10 km (rd.) W Chazuta, 6°37.157’S, 76°10.905’W, light, loc. 12, 10.I. 2009, leg. T. MALM & K. A. JOHANSON (NRM).

Etymology – Lemeza, from “lemez”, plate in Hungarian, refers to the subquadratic plate-like form of tergite X in dorsal view.

sp. n. (Figs 189–192)

Diagnosis – This species has an unusual short and high segment X, shell-like in dorsal view, with some similarity to S.(R.) abrupta FLINT, 1974 from Surinam, from which it is distinguished by the pair of deep shell-like concavities on dorsum X and by the phallic apex with 2 lateral processes and 2 lateral lobes.

Description – Male. Pale brown, forewing length 5 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth darker and zigzag bordered; nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. Male genitalia. Segment IX with stalked ventrum, anterodorsum indistinct; dorsal keel wide-rounded; intersegmental profile between segments IX and X low. Segment X short and high; vestigial paraproct forming heavily sclerotised pair of ventrolateral straps

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)271 directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopods, coxopodite high with very low basal part in lateral view; harpagones long slender slightly bending mesad and ventrad. Phallotheca with vertical inflated basal portion curving gradually to stem; phallic apex with more sclerotised dorsal part and complex ventral part; this ventral part composed of a pair of rounded vertical plates and a pair of short process tipped with short teeth; sclerotised section of ejaculatory duct short.

Material examined – Holotype, male: : Otun Quinbaya National Park, La Suiza, 1900 m, 13.III.2011, leg. J. OLÁH Jr. (OPC). Paratypes: same data as holotype (8 males, OPC). : Dept. Pasco, Yanachaga Chemilien NP, INRENA Refugio El Cedro, S10°32.717, W75°21.492, 2460 m, 30.I.2003, leg. A. KUN &B.BENEDEK, HNHM expedition, (2 males, HNHM). Pasco Reg., Yanachaga-Chemillen N.P., side river to Rio Huan- cabamba, N end of park, along Oxabamba-Pozuzo rd., 10°11.133’S, 75°34.106’W, light, loc.01 31.XII.2008, leg. T. MALM & K. A. JOHANSON (11 males, NRM, 10 males OPC).

Etymology – Medena, coined from “medence”, basin in Hungarian, refers to the pair of deep concavities on segment X.

sp. n. (Figs 193–197)

Diagnosis – This new species is a member of the S.(S.) nigripennis species group, and related to S.(S.) turrialbana FLINT, 1974 from Costa Rica, but easily distinguished by the forewing having no 2 transverse white bands; by the V-shaped straight ventroapical pair of processes on the phallotheca, not hooked sharply laterad in right angle.

Description – Male. Dark, almost black forewing length 4 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands without long anterolateral filament, but with 2 pairs of internal reticulate sacs in subapical abdominal segments. Forewing membrane dark homogenous with lighter area on pterostigma and arculus. Male genitalia. Segment IX produced anterad; intersegmental profile between segments IX and X almost indistinct. Segment X short, lateral lobes with small excision; cerci small represented by a few setae; vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin, functioning as sclerotised phallic guides for phallic organ. Basal segment of gonopods, coxopodite slender, expanded slightly apicad; harpagones long slender parallel-sided slightly clavate in lateral view. Phallotheca with apical processes straight V-shaped.

Material examined – Holotype, male: : Minca, 650 m, 24.III.2011, light, leg. J. OLÁH Jr. (OPC).

Etymology – Minca from the type locality.

Annls hist.-nat. Mus. natn. hung. 104, 2012 272 J. Oláh & K. A. Johanson

190

189

192 191

Smicridea (Rhyacophylax) medena sp. n., holotype, male genitalia: 189 = lateral view; 190 = left gonopod, ventral; 191 = dorsal; 192 = phallus lateral

193 194

196

195 197

Smicridea (Smicridea) mincana sp. n., holotype, male genitalia: 193 = lateral view; 194 = left gonopod, ventral; 195 = dorsal; 196 = phallus lateral; 197 = phalus ventral

Smicridea (Smicridea) mucronata FLINT et DENNING, 1989

Material examined – : Nequén, Parque Nacional Lanin, small stream crossing road RP61, 5 km E Arroyo Raquithué, 25 m upstream road, 908 m, 39°44’52.9”S, 71°27’ 52.4” W, loc Ar5, light trap 12.II.2010, leg. M. ESPELAND (1 male, NRM). : Region del Araucania (IX), PN, Nahuelbuta, camp site, Estero Cabreria, crosspoint between streams draining Mts Pichinahuel and Pichimanquemáhuida, 37°49.647’S, 73°00.691’W, 1100 m (Loc #07), Malaise trap 3–4.I.2006, leg. K. A. JOHANSON (2 males, NRM, 2 males, OPC).

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)273

Smicridea (Rhyacophylax) murina MCLACHLAN, 1871 (Figs 198–201, 202–206)

Material examined – : Jardin, West Andes, 1750 m, 17.III.2011, light, leg. J. OLÁH Jr. (1 male, OPC). : Western Andean Slope, Alambi, 24.IX.2010, light, leg. J. OLÁH Jr. (1 male, OPC). Amasonian Lowland, Gareno Lodge, 18.XI.2006, light, leg. J. OLÁH Jr. (1 male, OPC). Amasonia, Gareno Lodge, Eastern Andean Slope, 13–14.VIII. 2011, light, leg. J. OLÁH Jr. (6 males, OPC). : Dept. Lima, Pacaran, Prov. Canete, River Chillon Obrajillo, 877 m, 76°02’60”W, 12°52’05”S, 16–17.III.2006, light trap, leg. FAVIOLA MONTES CARLOS &JUAN CHAVEZ LOPEZ (58 males, 93 females, NRM). Dept. Lima, Catahuasi District, Yruyos Province, River Tupe, 1581 m, 75°53’3”W, 12°47’56”S, 20–21.III. 2006, light trap, leg. FAVIOLA MONTES CARLOS &JUAN CHAVEZ LOPEZ (25 males, NRM). Dept. Lima, Catamuasi District, Yauyos Province, River Canete, Pob. Cacacho, El Anbo, 1489 m, 75°54’05”W, 12°48’05”S, 22–23.III.2006, light trap, leg. FAVIOLA MONTES CARLOS &JUAN CHAVEZ LOPEZ (11 males, 32 females, NRM). San Martin Prov., Rio Negro, 37 km (rd.) W Moyobamba, near Olmos-Tarapoto rd., 6°00.278’S, 77°15.437’W, light, loc. 05, 6.I.2009, leg. K. A. JOHANSON & T. MALM (40 males, NRM, 8 males, OPC).

Remarks – The strengthening frame of the combined paraproct and antecostal suture complex less developed on the Alambi Ecuadorian specimens, especially the dorsal section of antecostal suture is abbreviated (Fig. 202). The dorsoapical cornuted processes less developed on Gareno Ecuadorian specimen (Fig. 198).

199

198

200 201

Smicridea (Rhyacophylax) murina MCLACHLAN, 1871, male genitalia: 198 = lateral view; 199 = left gonopod, ventral; 200 = dorsal; 201 = phallus lateral. Gareno Ecua- dorian specimen

Annls hist.-nat. Mus. natn. hung. 104, 2012 274 J. Oláh & K. A. Johanson

203

202

205 204

206

Smicridea (Rhyacophylax) murina MCLACHLAN, 1871, male genitalia: 202 = lateral view; 203 = left gonopod, ventral; 204 = dorsal; 205 = phallus lateral; 206 = phallus apex, ventral. Alambi Ecuadorian specimen

sp. n. (Figs 207–211)

Diagnosis – This species belongs to the S.(R.) signata group, and is closest to S. (R.) bidactyla FLINT et REYES-ARRUNATEGUI, 1991 from Ecuador, from which it is distinguished by having different structure of strengthening frame on segments IX and X; a single ventrolateral paraproctal bifid spike on segment X, not bidactylate; no dorsal and ventral subapical spines clusters present on phallic organ.

Description – Male. Pale brown, forewing length 4.5 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: white transverse band at apical fourth; chord (anastomosis) of crossveins s, r-m, m, m-cu and cu darker. Male genitalia. Segment IX subtriangular with vertical stalk, anterodorsum indistinct; dorsal keel rounded; intersegmental profile between segments IX and X indistinct. Segment X short, lateral lobes with pointed tip with a subapical small ventral bifid spike; several short spicules on mesal surface of lateral lobes; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral straps directed downward from apical margin, fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame; 2 additional small suture branches developed. Basal segment of gonopods, coxopodite stalked and straight and reaching to tip of segment X; harpagones slender parallel-sided slightly bending mesad. Phallotheca with low vertical inflated basal portion and apex with ventral tongue-like process accompanied by short lateral teeth-tipped processes.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)275

Material examined – Holotype, male: : Amasonian Lowland, Gareno, near Puerto Napo, 400 m, 14.VIII.2011, light, leg. J. OLÁH Jr. (OPC). Paratypes: : Loreto Prov., stream crossing rd. Jurimaguas-Maniche, 2 km (rd.) W Yurimaguas, 5°53.934’S, 76°07.916’W, light, loc. 13, 11.I.2009, leg. K. A. JOHANSON &T.MALM (1 male, NRM). San Martin Prov., Rio Huallaga tributary, small river passing Chazuta, 6°34.665’S, 76°08.209’ W, light, loc. 11, 10.I.2009, leg. T. MALM &K.A.JOHANSON (4 males, NRM, 2 males, OPC). San Martin Prov., stream crossing Juan Guerra-Chazuta rd., 10 km (rd.) W Chazu- ta, 6°37.157’S, 76°10.905’W, light, loc. 12, 10.I.2009, leg. T. MALM &K.A.JOHANSON (19 males, NRM). San Martin Prov., creek crossing rd. Juan Guerra-Chazuta, 14 km (rd.) E Colombia Bridge, 6°35.594’S, 76°13.172’W, light, loc. 09, 9.I.2009, leg. K. A. JOHANSON & T. MALM (2 males, NRM).

Etymology – Nemtompa, from “nem tompa”, not blunt in Hungarian, refers to pointed tips of the lateral lobes on segment X.

207 208

211

210 209

Smicridea (Rhyacophylax) nemtompa sp. n., holotype, male genitalia: 207 = lateral view; 208 = left gonopod, ventral; 209 = dorsal; 210 = phallus lateral; 211 = phallus apex, ventral

sp. n. (Figs 212–216)

Diagnosis – This species, with apicodorsal phallic lobes, belongs to the S.(R.) murina species group, and is closest to S.(R.) murina MCLACHLAN, 1871, from which it differs by having lateral lobes on segment X differently shaped in dorsal view, their lateral margin convex, not concave; dorsum of phallotheca concave in lateral view, not convex; dorsoapical processes on phallic apex small.

Annls hist.-nat. Mus. natn. hung. 104, 2012 276 J. Oláh & K. A. Johanson

Description – Male. Pale brown, forewing length 4.8 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: white transverse band at apical fourth; chord (anastomosis) with darker patches around crossveins s, r-m, m, m-cu and cu. Male genitalia. Segment IX subtriangular with stalked ventrum; anterodorsum indistinct; dorsal keel rounded triangular; intersegmental profile between segments IX and X acute angled. Lateral lobes on segment X with mesad directed tip; cerci rounded ovoid; vestigial paraproct forming heavily sclerotised pair of ventrolateral straps directed downward from apical margin and fused with antecostal surure of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite straight, parallel- sided and reaching over tip of segment X; harpagones slender parallel-sided slightly bending mesad. Phallotheca with high vertical inflated basal portion; apex with a pair of distinct dorosapical short processes.

213

212

216 214

215

Smicridea (Rhyacophylax) parany sp. n., holotype, male genitalia: 212 = lateral view; 213 = left gonopod, ventral; 214 = dorsal; 215 = phallus lateral; 216 = phallus apex, ventral

Material examined – Holotype, male: : Amazonas Prov., Rio Utcabamba, Bajra Grande, at Rio Hotel, 5°45.824’S, 78°25.414’W, light, loc. 14, 12.I.2009, leg. T. MALM &K. A. JOHANSON (NRM). Paratypes: same data as holotype (2 males, NRM, 2 males, OPC). San Martin Prov., Rio Mayo, 11 km (rd.) E Mayobamba, 6°04.989’S, 76°53.065’W, light, loc. 06, 6.I.2009, leg. T. MALM & K. A. JOHANSON (4 males, NRM, 2 males, OPC).

Etymology – Parany, from “parány”, tiny in Hungarian, refers to the small dorsoapical phallic processes.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)277

Smicridea (Smicridea) penai FLINT et DENNING, 1989

Material examined – : Region de los Lagos (X), Isla Grande de Chiloé, Rio Meli- lebú, along road between Tebuhueico and Hullinco, 5.4 km S crosspoint to Curaco, river, 100 m upstream bridge, ca. 42°43.089’S, 73°53.797’W, 43 m (Loc #15), light trap, leg. K. A. JOHANSON (8 males, NRM, 5 males, OPC). Region de los Lagos (X), Isla Grande de Chiloé, stream crossing “Historico” road 1 km W Dalcahue, 42°22.734’S, 73°40.447’W, 34 m (Loc #17), light trap, leg. K. A. JOHANSON (2 males, NRM).

Smicridea (Rhyacophylax) peruana (MARTYNOV, 1912) (Figs 217–219)

Material examined – : Huanuco, stream at Chinchao, 2200 m, 76°00’W, 9°66’S, 18–26.VI.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (40 males, 700 females NRM, 21 males, 50 females OPC). Huanuco, stream at Carpish, 2500 m, 76°09’W, 9°40’S, 28.VI–3. VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (22 males, 400 females NRM, 12 males, 50 females, OPC). Dept. Pasco, Yanachaga Chemilien NP, INRENA Refugio El Cedro, S10°32.717, W75°21.492, 2460 m, 30.I.2003, leg A. KUN &B.BENEDEK, HNHM expedition, (40 males, HNHM). Pasco Reg., Yanachaga-Chemillen NP, side river to Rio Huancabamba, N end of park, along Oxabamba-Pozuzo rd., 10°11.133’ S, 75°34.106’W, light, loc. 01 31.XII.2008, leg. T. MALM &K.A.JOHANSON (15 males, NRM, 10 males OPC).

Smicridea (Rhyacophylax) pseudoradula FLINT, 1991

Material examined – : San Martin Prov., Rio Huallaga tributary, small river passing Chazuta, 6°34.665’S, 76°08.209’W, light, loc. 11, 10.I.2009, leg. T. MALM &K.A. JOHANSON (7 males, NRM, 6 males, OPC).

Smicridea (Smicridea) pucara FLINT et DENNING, 1989

Material examined – : Nequén, Parque Nacional Lanin, Small stream crossing road RP61, 5 km E Arroyo Raquithué, 25 m upstream road, 908 m, 39°44’52.9”S, 71°27’ 52.4”W, loc Ar5, light trap 12.II.2010, leg. M. ESPELAND (2 males, NRM, 2 males, OPC).

Smicridea (Smicridea) rarus BUENO-SORIA et MÁRQUEZ-MAYAUDON, 1979

Annls hist.-nat. Mus. natn. hung. 104, 2012 278 J. Oláh & K. A. Johanson

217

219 218

Smicridea (Rhyacophylax) peruana (MARTYNOV, 1912), male genitalia: 217 = lateral view; 218 = dorsal; 219 = phallus lateral

sp. n. (Figs 220–225)

Diagnosis – This species belongs to S. (R.) signata group, and is closest to S. (R.) lobata (ULMER, 1909) from Venezuela and S. (R.) nemtompa sp. n. from Ecuador and Peru, but easily distinguished by having lateral wing-shaped processes on the middle of the phallotheca.

Description – Male. Pale brown, forewing length 4.5 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: white transverse band at apical fourth; chord (anastomosis) of crossveins s, r-m, m, m-cu and cu darker. Male genitalia. Segment IX subtriangular with high vertical stalk, anterodorsum indistinct; dorsal keel rounded; intersegmental profile between segments IX and X right angled. Segment X medium long, lateral lobes with blunt tip with a subapical small bifid ventral paraproctal spike; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral straps directed downward from subapical spike, fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite stalked and straight and reaching over tip of segment X; harpagones slender parallel-sided slightly bending mesad. Phallotheca with low vertical inflated basal portion and wingshaped spiny lateral processes midway; phallic apex with dorsal membranous bilobed plate, ventral tongue-like process accompanied by short lateral teeth-tipped processes; eversible phallotremal and endothecal complex tele- scopic with a pair of short spines.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)279

221 220

223 225

222 224

Smicridea (Rhyacophylax) repula sp. n., holotype, male genitalia: 220 = lateral view; 221 = left gonopod, ventral; 222 = dorsal; 223 = phallus lateral, erected; 224 = phallus lateral, not erected; 225 = lateral process midway on the phallic organ

Material examined – Holotype, male: : State of Veracruz, Los Tuxtlas area, Rio la Palma, near to the Estacion the Biologia Los Tuxtlas, N 18°33.680’, W 095°02.943’, 30 mao, 26.VI.2006, light trap, leg. M. ESPELAND &T.MALM (NRM). Paratypes: same data as holotype (24 males, NRM, 3 males, OPC). State of Veracruz, Fortin de las Flores, Metlac River, under bridge of R 150, 900 mao, 24.VI.2006, light trap, leg. M. ESPELAND & T. MALM (1 male, NRM). State of Veracruz, Los Tuxtlas area, Rio Maquinas, near to the Estacion the Biologia Los Tuxtlas, N 18°37.392’, W 095°05.418’, 9 mao, 27.VI.2006, light trap, leg. M. ESPELAND & T. MALM (11 males, NRM).

Etymology – Repula, from “repülõ”, flying in Hungarian, refers to wing-shaped spiny lateral processes midway on the phallotheca.

sp. n. (Figs 226–230)

Diagnosis – S. (R.) radula FLINT, 1974, S. (R.) pseudoradula FLINT, 1991, S. (R.) kapara sp. n. and this new species are very close. The new species differs from the other three species by having dorsoapical lobe on phallic apex narrowing apicad and the radulalike ventrolateral lobes inverted mesad and lost its free lobelike form.

Description – Male. Small pale brown animal, forewing length 4 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long

Annls hist.-nat. Mus. natn. hung. 104, 2012 280 J. Oláh & K. A. Johanson anterolateral filament. Forewing membrane without distinct patterne, except darker apical region and crossveins more pigmented. Male genitalia. Segment IX with anterolateral margin produced as a narrow, slightly upturned lobe; sternum with a short, pointed process between gonopod bases; intersegmental profile between segments IX and X right angled. Segment X short, lateral lobes short and blunt; cerci rounded ovoid, located apical. Vestigial paraproct forming heavily sclerotised pair of ventrolateral internal straps with apical apex projecting with double teeth; paraproctal straps fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite S-forming and reaching over tip of segment X; harpagones slender. Phallotheca with inflated vertical basal portion and apex with narrowing dorsal lobe and with a pair of mesad inverted ventrolateral radulalike rasps.

227 226

229

228 230

Smicridea (Rhyacophylax) resela sp. n., holotype, male genitalia: 226 = lateral view; 227 = left gonopod, ventral; 228 = dorsal; 229 = phallus lateral; 230 = phallus apex, ventral

Material examined – Holotype, male: : San Martin Prov., creek crossing rd. Ta- rapoto-Yurimaguas, ca. 30 km (rd.) NE Tarapoto, 6°24.904’S, 76°18.756’W, light, loc. 08, 8.I.2009, leg. K. A. JOHANSON &T.MALM (NRM). Paratype: same data as holotype (1 male, OPC).

Etymology – Resela, from “reszelõ”, rasp in Hungarian, refers to the dentate radula-like ventrolateral lobes on the phallic apex.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)281

sp. n. (Figs 231–235)

Diagnosis – This species is closely related to S.(R.) biserrulata FLINT, 1991 from Co- lombia, but differs in dorsal view by the mesally angled tip of the lateral lobes of segment X, not narrowly rounded; the apex of phallic organ with a pair of dorsolateral short process tipped with short spines, not with a series of points without any process.

Description – Male. Pale brown, forewing length 6.7 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth darker and zigzag bordered; nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. Male genitalia. Segment IX subtriangular, anterodorsum indistinct; dorsal keel triangular tipped with microtrichiae; intersegmental profile between segments IX and X almost right angled with a small membraneous process below tip of dorsal keel; ventromesally with a posterior long triangular lobe. Segment X long, lateral lobes with mesally angled tip with minute pointed process; cerci longitudinally ovoid, located little nearer to basis; vestigial paraproct forming heavily sclerotised pair of internal straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopods, coxopodite high with very low basal fourth in lateral view; harpagones long slender parallel-sided slightly bending mesad and ventrad. Phallotheca with vertical inflated basal portion and apex with asperites of ventrolateral row of small spine points separated by ventromesal dark keel; a pair of short subapical processes dorsolaterally tipped with 3–4 small spines; tip composed of indistinct ventral and dorsal lobes plus very tip of ejaculatory duct projecting free as a continuation of its internal more pigmented line, not angled at midlength.

Material examined – Holotype, male: : Mindo, Septimo Paraiso, 1600 m, 5.VIII.2011, light, leg. J. OLÁH Jr. (OPC). Paratypes: same data as holotype (1 male, 2 females, OPC). Mindo, Septimo Paraiso, 1600 m, 30.IX.2010, light, leg. J. OLÁH Jr. (4 males, 3 females, OPC). Mindo, Septimo Paraiso, 1600 m, IX.2008, light, leg. J. OLÁH Jr. (1 male, OPC). Wild Sumaco, near Pacto Sumaco, 7–8.X.2010, light, leg. J. OLÁH Jr. (2 males, 7 females, OPC). Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 24.XI.2007, light, leg. J. OLÁH Jr. (4 males, OPC). Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 15.X.2010, light, leg. J. OLÁH Jr. (1 male, OPC). Tina- landia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 19.VIII.2011, light, leg.J.OLÁH Jr.(2males,OPC). : Otun Quinbaya National Park, La Suiza, 1900 m, 13.III.2011, leg. J. OLÁH Jr. (1male, OPC). Jardin, West Andes, 1750 m, 17.III.2011, light, leg. J. OLÁH Jr. (3 males, 6 females, OPC).

Etymology – Sarkoska, from “sarkoska”, diminutive of angled in Hungarian, refers to mesally angled tips of the lateral lobes on segment X.

Annls hist.-nat. Mus. natn. hung. 104, 2012 282 J. Oláh & K. A. Johanson

232

231

235

233 234

Smicridea (Rhyacophylax) sarkoska sp. n., holotype, male genitalia: 231 = lateral view; 232 = left gonopod, ventral; 233 = dorsal; 234 = phallus lateral; 235 = phallus apex, ventral

sp. n. (Figs 236–240)

Diagnosis – This species belongs to a group of species having robust and short coxopodite: S. breviuncata FLINT, 1974, S. conjuncta FLINT, 1991, S. nigerrima FLINT, 1983, S. prorigera FLINT, 1991, S. ruginasa FLINT, 1991, S. saucia MCLACHLAN, 1871, and S. urra FLINT, 1991. It is most similar to S. saucia from Peru, from which it differs by having differently shaped segment X, phallic apex with different sclerites and harpago distinctly mesad curving falciform.

Description – Male. Almost uniformly fuscous with unmarked wings, forewing length 6.3 mm. Maxillary palp formula 1–2–3–4–5. Tibial spurs 144. Cuticular structure of sternum V glands without long anterolateral filament, but with 2 pairs of internal reticulate sacs in subapical abdominal segments. Male genitalia. Segment IX produced middle anterad; intersegmental profile between segments IX and X very sharp angled. Segment X with rather unusual specific shape; cerci small located between lateral lobes and dorsal keels; vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin. Basal segment of gonopods, coxopodite robust and short, harpago falciform. Phallotheca with rounded apex composed of lateral shells and middle spine.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)283

236 237

238

239 240

Smicridea (Smicridea) sarla sp. n., holotype, male genitalia: 236 = lateral view; 237 = left gonopod, ventral; 238 = dorsal; 239 = phallus lateral; 240 = phallus apex, ventral

Material examined – Holotype, male: : Huanuco, stream at Carpish, 2500 m, 76°09’W, 9°40’S, 28.VI–3.VII.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (1 male, NRM). Paratypes: Same data as holotype (6 males, 17 females, NRM, 4 males, 10 females, OPC). Huanuco, stream at Chinchao, 2200 m, 76o00’W, 9o66’S, 18–26.VI.2005, Malaise trap, leg. FAVIOLA MONTES CARLOS (19 males, 12 females, NRM).

Etymology – Sarla, from “sarlós”, sickle shaped in Hungarian, refers to the sickle–like falciform harpago.

sp. n. (Figs 241–246)

Diagnosis – This species, with distal end of phallotheca simple and without pronounced dorsal, lateral or ventral processes, belongs to the S.(R.) dispar species group, and having phallotremal sclerites elongated scissor-shaped belongs to the S.(R) appendiculata species complex. It is closest to S.(R.) bicornuta ALBINO,PES et HAMADA, 2011, from which it differs by having dorsal keel of segment IX narrow triangular in dorsal view, not broad; phallotheca robust in lateral view, not slender; endothecal hornlike processes and phallotremal sclerites differently shaped.

Description – Male. Pale brown, forewing length 5 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long antero-

Annls hist.-nat. Mus. natn. hung. 104, 2012 284 J. Oláh & K. A. Johanson lateral filament. Forewing membrane patterned: apical fourth slightly darker with zigzag margin; chord (anastomosis) of crossveins s, r-m, m, m-cu and cu darker. Male genitalia. Segment IX subtriangular with stalked ventrum; anterodorsum indistinct; dorsal keel narrow triangular in dorsal view; intersegmental profile between segments IX and X acute angled. Lateral lobes on segment X with blunt narrowing apex in dorsal view; cerci rounded ovoid, located middle; vestigial paraproct forming sclerotised pair of ventrolateral straps directed downward from apical margin and fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame; Basal segment of gonopods, coxopodite straight, almost parallel-sided and reaching to tip of segment X; harpagones slender parallel-sided slightly bending mesad. Phallotheca robust; apex without processes or lobes, long endophallus and elongated phallotremal scissorlike sclerites, endotheca armed with hornlike processes.

Etymology – Sarvaka, from “szarvak”, horns in Hungarian, refers to the pair of horns in the endotheca, well visible especially when endotheca is evaginated.

242

241

246

243 244 245

Smicridea (Rhyacophylax) sarvaka sp. n., holotype, male genitalia: 241 = lateral view; 242 = left gonopod, ventral; 243 = dorsal; 244 = phallus lateral; 245 = phallus lateral, erected; 246 = phallus apex, ventral

Smicridea (Rhyacophylax) scutellaria FLINT, 1974

Material examined – : Maripasoula, Maroni River, Damason campo, Village, 4°35.112’N, 54°24.799’W, 38 m, 7.II.2007, FRG 13, leg. N. JÖNSSON (20 males,

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)285

7 females, NRM, 20 males, 10 females, OPC). Maripasoula, Lawa River: Maripasoula, 83 mao, 3°37.959’N, 54°1.426’W, 9.II.2007,light trap, FRG 15, leg. N. JÖNSSON (20 males, 7 females, NRM, 47 males, 10 females, OPC).

sp. n. (Figs 247–250)

Diagnosis – This species belongs to S.(R.) peruana species group, and is very closely related to S. (R.) acuminata FLINT, 1974 from Costa Rica, but distinguished in dorsal view by having the stalked ventrum of segment IX high in lateral view, not low; lateral lobes on segment X straight, not curving mesad; apex of phallic organ with a spine cluster of 4 spines, not 2 spines only.

Description – Male. Pale brown, forewing length 7 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth darker and zigzag bordered; nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. Male genitalia. Segment IX with high staked ventrum in lateral view, anterodorsum indistinct; dorsal keel triangular covered with microtrichiae; intersegmental profile between segments IX and X sharp angled; ventromesally with a posterior triangular lobe. Seg- ment X long, lateral lobes straight long triangular; cerci longitudinally ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral straps on segment X directed downward from apical margin and fused to suture on ventrum IX, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopods, coxopodite high with low basal fourth in lateral view; harpagones long slender parallel-sided slightly bending mesad and ventrad. Phallotheca with vertical inflated basal portion turning gradually to stem, apex with asperites of dorsolateral patch and ventromedian row of spicules; apicoventrum bilobed, apicodorsum monolobed; internal spine cluster visible having 4 free tips of spines.

Material examined – Holotype, male: : Western Andean Slope, Alambi, 24.IX.2010, light, leg. J. OLÁH Jr. (1 male, OPC). Paratypes: same data as holotype (2 males, OPC). Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 19.VIII.2011, light, leg. J. OLÁH Jr. (3 males, OPC). : San Martin Prov., creek crossing rd. Tarapoto-Yurimaguas, ca. 30 km (rd.) NE Tarapoto, 6°24.904’S, 76°18.756’W, light, loc. 08, 8.I.2009, leg. K. A. JOHANSON &T.MALM (2 males, NRM). San Martin Prov., Rio Negro, 37 km (rd.) W Moyobamba, near Olmos-Tarapoto rd., 6°00.278’S, 77°15.437’W, light, loc. 05, 6.I.2009, leg. K. A. JOHANSON & T. MALM (2 males, NRM).

Etymology – Sudara, from “sudár”, straight in Hungarian, refers to straight and slender lateral lobes on segment X.

Annls hist.-nat. Mus. natn. hung. 104, 2012 286 J. Oláh & K. A. Johanson

247

248

249 250

Smicridea (Rhyacophylax) sudara sp. n., holotype, male genitalia: 247 = lateral view; 248 = dorsal; 249 = phallus lateral; 250 = phallus apex, ventral

sp. n. (Figs 251–255)

Diagnosis – This species belongs to the S.(R.) signata group, and is close to S.(R.) hajla sp. n. from Ecuador, but differs by having apical fourth of forewing membrane darker, not with white transverse band at apical fourth; lateral lobes on segment X long and straight in dorsal view, not short, tapering and mesad curving; phallic apex differently formed.

Description – Male. Pale brown, forewing length 4.8 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth darker and zigzag bordered; nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. Male genitalia. Segment IX subtriangular with vertical stalk, anterodorsum indistinct; dorsal keel rounded, microtrichous; intersegmental profile between segments IX and X obtuse angled. Segment X short, lateral lobes with straight; a pair of serrated and spiny apicoventral margin present; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of lateroventral straps directed downward from apical margin, functioning as sclerotised phallic guides encircling above phallic organ; paraproctal straps confused with antecostal suture of segment IX forming together an Y-shaped strengthening frame, a rather well-sclerotised strap forming a dorsal guide on phallic organ. Basal segment of gonopods, coxopodite stalked and straight, and reaching to tip of segment X; harpagones slender parallel-sided slightly bending mesad. Phallotheca with low vertical inflated basal portion and apex with ventral lobe; phallic dorsum with fused paraproctal attachment.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)287

252

251

255

253 254

Smicridea (Rhyacophylax) tavola sp. n., holotype, male genitalia: 251 = lateral view; 252 = left gonopod, ventral; 253 = dorsal; 254 = phallus lateral; 255 = phallus apex, ventral

Material examined – Holotype, male: : Amasonian Lowland, Gareno, near Puerto Napo, 400 m, 14.VIII.2011, light, leg. J. OLÁH Jr. (OPC). Paratypes: same data as holotype (2 males, OPC).

Etymology – Tavola, from “távol”, far in Hungarian, refers to the lateral lobes located far from each others on segment X.

sp. n. (Figs 256–260)

Diagnosis – This species, with apicodorsal phallic lobes, belongs to the S.(R.) murina species group, and is most similar to S.(R.) furesa sp. n., and differs by having lateral processes on segment X elongated with mesad turning apices; dorsoapical horn on phallic apex intermediate without mesal hump in ventral view; the ventrum of phallic apex without serrulate lines.

Annls hist.-nat. Mus. natn. hung. 104, 2012 288 J. Oláh & K. A. Johanson angled. Segment X long, lateral lobes with mesad turning tip; cerci rounded ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of ventrolateral straps directed downward from apical margin and fused with antecostal surure of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite straight and reaching before tip of segment X; harpagones slender parallel-sided slightly capitate. Phallotheca with vertical inflated basal portion almost as high as length of stem; apex with distinct dorsoapical thick horns and sclerotised apical half of ejaculatory duct, ending in phallotermal sclerites like a pair of spines. 257

256

258 259

260

Smicridea (Rhyacophylax) tina sp. n., holotype, male genitalia: 256 = lateral view; 257 = left gonopod, ventral; 258 = dorsal; 259 = phallus lateral; 260 = phallus apex, ventral

Material examined – Holotype, male: : Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 24.XI.2007, light, leg. J. OLÁH Jr. (OPC). Para- types: : Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 15.X.2010, light, leg. J. OLÁH Jr. (1 male, OPC). : Amazonas Prov., Rio Utcabamba, Bajra Grande, at Rio Hotel, 5°45.824’S, 78°25.414’W, light, loc. 14, 12.I.2009, leg. T. MALM &K.A.JOHANSON (1 male, NRM, 1 male OPC). San Martin Prov., Rio Mayo, 11 km (rd.) E Mayobamba, 6°04.989’S, 76°53.065’W, light, loc. 06, 6.I.2009, leg. T. MALM &K.A. JOHANSON (2 males, NRM).

Etymology – Tina, name derived from the type locality.

Smicridea (Rhyacophylax) titschacki FLINT, 1975

Material examined – : San Martin Prov., Rio Huallaga, at Pumarihri Huallaga Lodge, between Juan Guerra and Chazuta, 14 km (rd) W Chazuta, 6°36.643’S, 76°12.555’

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)289

W, light, loc. 10, 9.I.2009, leg. K. A. JOHANSON &T.MALM (9 males NRM, 7 males OPC). Amazonas Prov., Rio Utcabamba, Bajra Grande, at Rio Hotel, 5°45.824’S, 78°25.414’W, light, loc. 14, 12.I.2009, leg. T. MALM &K.A.JOHANSON (1 male, NRM). San Martin Prov., Rio Negro, 37 km (rd.) W Moyobamba, near Olmos-Tarapoto rd., 6°00.278’S, 77°15.437’ W, light, loc. 05, 6.I.2009, leg. K. A. JOHANSON &T.MALM (1 male, OPC). San Martin Prov., Rio Mayo, 11 km (rd.) E Mayobamba, 6°04.989’S, 76°53.065’W, light, loc. 06, 6.I.2009, leg. T. MALM & K. A. JOHANSON (2 males, OPC).

sp. n. (Figs 261–265)

Diagnosis – This species belongs to the S.(S.) fasciatella species group, and is close to S.(S.) truncata FLINT, 1974 from Surinam, from which it is distinguished by having segment IX low, not high; apicodorsal excision on segment IX rectangular in lateral view, not obtuse angled; lateral processes of segment X with narrowing apices, not blunt; basal vertical section of phallotheca high, not low, phallic head capitate.

262 261

263 264 265

Smicridea (Smicridea) torpa sp. n., holotype, male genitalia: 261 = lateral view; 262 = left gonopod, ventral; 263 = dorsal; 264 = phallus lateral; 265 = phallus apex, ventral

Description – Male. Dark, almost black without any membrane pattern except hyaline line along stem M; forewing length 5 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands with anterolateral filament as long as segment Y; with 2 pairs of internal reticulate sacs present in subapical abdominal segments.

Annls hist.-nat. Mus. natn. hung. 104, 2012 290 J. Oláh & K. A. Johanson

Male genitalia. Segment IX straight vertical anterad with rectangular apicodorsal excision in lateral view; intersegmental profile between segments IX and X absent. Segment X enlarged, lateral processes tapering in dorsal view; cerci large located laterally subapicad; vestigial paraproct forming sclerotised pair of lateroventral straps directed downward from apical margin, its basal part more developed. Basal segment of gonopods, coxopodite high, low in very basad; harpagones truncate in lateral view, mesal pointed in ventral view. Phallotheca with high basal section and with capitate membranous apex.

Material examined – Holotype, male: : Approuaguekaw, Kaw Mt, 104 mao, 4°33.035’N, 52°11.661’W, Malaise trap, 23.I–7.II.2007, FRG MF3, leg. N. JÖNSSON (NRM).

Etymology – Torpa, coined from “törpe”, pygmy in Hungarian, refers to the low segment IX.

sp. n. (Figs 266–270)

Diagnosis – This species belongs to the S.(S.) nigripennis group, and is closest to S. (S.) mirnae ALMEIDA et FLINT, 2002 from Brazil, from which it is distinguished by having segment IX with anterodorsal excision less developed; lateral lobes of segment X broad, not narrow; endotheca with a pair of long spines, not with seven small spines.

Description – Male. Dark brown, forewing length 5 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands with long anterolateral filament, 2 pairs of internal reticulate sacs in subapical abdominal segments present. Fore- wing membrane dark homogenous with hyaline window pattern only present along stem M1 and on Cu2 confluence at arculus. Male genitalia. Segment IX with basodorsal excision less developed. Segment X short, lateral processes broad truncate; cerci elongated embrassed by V-shaped vestigial paraproct of sclerotised pair of lateroventral straps directed downward from apical margin. Basal segment of gonopods, coxopodite stout; harpagones slightly capitate in lateral view and pointed in ventral view. Phallotheca with a pair of lateral processes and with an entire ventral lobe; endotheca with a pair long spines and a pair of rounding sclerites.

Material examined – Holotype, male: : Approuaguekaw, Kaw Mt, 216 m, 4°33.257’N, 52°11.920’W, Malaise trap, 19.I–4.II.2007, FRG MF 2, leg. N. JÖNSSON (NRM). Paratypes: same data as holotype (12 males, NRM, 5 males, OPC). Approuague- kaw, Kaw Mt, 104 m, 4°33.035’N, 52°11.661’W, Malaise trap, 4–12.II.2007, FRG MF3, leg. N. JÖNSSON (1 male, NRM).

Etymology – Vagotta, coined from “vágott”, truncate in Hungarian, refers to the broad- ly truncate lateral lobes on segment X.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)291

267

266

268

269 270

Smicridea (Smicridea) vagotta sp. n., holotype, male genitalia: 266 = lateral view; 267 = left gonopod, ventral; 268 = dorsal; 269 = phallus lateral; 270 = phallus ventral

sp. n. (Figs 271–275)

Diagnosis – This species belongs to the S.(R.) radula species complex and is closest to S.(R.) kapara sp. n., but differs by having phallotheca S-formed, not with straight stem; dorsal lobe on the phallic apex simply rounded, not stalked and scaled, not bare.

Description – Male. Small pale brown animal, forewing length 4 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane with distinct pattern, exhibiting dark apical region and crossveins. Male genitalia. Segment IX with anterolateral margin produced as a narrow, slightly upturned lobe; sternum with a narrow, pointed process between gonopod bases; intersegmental profile between segments IX and X right angled. Segment X short, lateral lobes short and blunt; cerci rounded ovoid, located apicad. Vestigial paraproct forming heavily sclerotised pair of ventrolateral internal straps with apical apex projecting with 2 teeth; paraproctal straps fused with antecostal suture of segment IX forming together an Y-shaped strengthening frame. Basal segment of gonopods, coxopodite straight and reaching over tip of segment X; harpagones slender. Phallotheca with inflated vertical basal portion and apex with rounded dorsal scaled lobe and a pair of ventrolateral lobes armed with a series of short, dark teeth recurving basad; sclerotised threadlike and less sclerotised tubelike structures of ejaculatory duct protruded.

Material examined – Holotype, male: : San Martin Prov., Rio Huallaga, at Pu- marihri Huallaga Lodge, between Juan Guerra and Chazuta, 14 km (rd) W Chazuta,

Annls hist.-nat. Mus. natn. hung. 104, 2012 292 J. Oláh & K. A. Johanson

6°36.643’S, 76°12.555’W, light, loc. 10, 9.I.2009, leg. K. A. JOHANSON &T.MALM (NRM). Paratypes: same data as holotype (3 males, NRM, 3 males, OPC). Amazonas Prov., Rio Utcabamba, Bajra Grande, at Rio Hotel, 5°45.824’S, 78°25.414’W, light, loc. 14, 12.I.2009, leg. K. A. JOHANSON &T.MALM (9 males, NRM). San Martin Prov., La Catarata de hua- shiyascu, 6°27.544’S, 76°18.192’W, light, loc. 07, 7.I.2009, leg. K. A. JOHANSON &T.MALM (1 male, NRM). San Martin Prov., Rio Mayo, 11 km (rd.) E Mayobamba, 6°04.989’ S, 76° 53.065’W, light, loc. 06, 6.I.2009, leg. T. MALM &K.A.JOHANSON (14 males, NRM, 10 males, OPC).

Etymology – Vakara, from “vakaró”, scraper in Hungarian, refers to the dentate radulalike ventrolateral lobes on the phallic apex.

272

271

275 273

274

Smicridea (Rhyacophylax) vakara sp. n., holotype, male genitalia: 271 = lateral view; 272 = left gonopod, ventral; 273 = dorsal; 274 = phallus lateral; 275 = phallus apex, ventral

sp. n. (Figs 276–279)

Diagnosis – This new species is a member of the S. (S.) nigripennis group, and most similar to S.(S.) inaequispina FLINT, 1974 from Surinam, but easily distinguished by having lateral profile of segment IX completely different; the single dorsomesal process of phallotheca well-developed, not indistinct; lateral processes on phallotheca present, not absent; endothecal spines stout, not slender.

Description – Male. Dark brown, forewing length 5 mm. Maxillary palp formula 1-2-3-4-5. Tibial spurs 144. Cuticular structure of sternum V glands with halfsegment long

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)293 anterolateral filament, 2 pairs of internal reticulate sacs in subapical abdominal segments present. Forewing membrane dark homogenous with hyaline window pattern only present along stem M1 and on Cu2 confluence at arculus. Male genitalia. Segment IX with long basodorsal excision in lateral view. Segment X short, lateral processes with small mesal angle; cerci elongated setose surface; vestigial paraproct of sclerotised pair of lateroventral straps directed downward from apical margin. Ba- sal segment of gonopods, coxopodite stout; harpagones long with blunt apices. Phallotheca with a single dorsomesal, a pair of lateral and a single ventromesal processes; endotheca with 2 pairs of stout and 1 pair of small slender spines.

277

276

278

279

Smicridea (Smicridea) vaskosa sp. n., holotype, male genitalia: 276 = lateral view; 277 = left gonopod, ventral; 278 = dorsal; 279 = phallus lateral

Material examined – Holotype, male: : Approuaguekaw, Kaw Mt, 4°33.235’N, 52°11.988’W, 225 mao, 20.I.2007 FRG 3, leg. N. JÖNSSON (NRM). Paratype: Approuaguekaw, Kaw Mt, 216 m, 4°33.257’N, 52°11.920’W, Malaise trap, 19.I–4.II.2007, FRG MF 2, leg. N. JÖNSSON (1 male, OPC).

Etymology – Vaskosa, from “vaskos”, stout in Hungarian, refers to the very stout spines in the endotheca.

sp. n. (Figs 280–282)

Diagnosis – This species belongs to the S.(R.) peruana species group, and is close to S. (R.) peruana (MARTYNOV, 1912), from which it differs by having forewing pattern more pronounced; tip of the lateral lobes on segment X mesad turning and narrowing lobe-like, not pointed spine-like; phallotheca slender in lateral view, not robust, dorsal asperite patch of small spinules less developed. S.(R.) peruana redescribed and redrawn from Argentina

Annls hist.-nat. Mus. natn. hung. 104, 2012 294 J. Oláh & K. A. Johanson and Bolivia (MARTIN &SGANGA 2011) as having pronounced forewing pattern, lobelike tip on lateral lobes and slender phallotheca probably represents S. (R.) vekona sp. n.

Description – Male. Pale brown, forewing length 8 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth darker and zigzag bordered; nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark; patternd very pronounced. Male genitalia. Segment IX subtriangular with high ventral stalk and ventroapical mesal triangular process; anterodorsum indistinct; dorsal keel long triangular covered with microtrichiae; intersegmental profile between segments IX and X acute angled. Segment X elongated, lateral lobes curving mesad with minute narrowing lobe; cerci longitudinally ovoid, located middle; vestigial paraproct forming heavily sclerotised pair of internal straps directed downward from apical margin. Basal segment of gonopods, coxopodite high with low basal fourth in lateral view; harpagones long slender, parallel-sided and slightly bending mesad and ventrad. Phallotheca thin, slender with vertical inflated basal portion right angled to arching horizontal stem; phallic apex with bifid dorsal process; dorsal asperite patch small ventral asperite row of small spinules long; phallotremal sclerites (internal sclerite) long and thin.

Material examined – Holotype, male: : Jujuy, PN, Calilegua, 13 km NW, visitor center, RP 83, 1263 m, 23°41.923’S, 64°52.342’W, 14.I.2008, leg. K. HILL,D.MARSHALL &M.MOULDS (NRM). Paratypes: same data as holotype (3 males, NRM, 2 males, OPC).

Etymology – Vekona, from “vékony”, thin, slender in Hungarian, refers to the very slender, thin phallotheca.

280

281

282

Smicridea (Rhyacophylax) vekona sp. n., holotype, male genitalia: 280 = lateral view; 281 = dorsal; 282 = phallus lateral

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)295

Smicridea (Rhyacophylax) ventridenticulata FLINT, 1991

Material examined – : Otun Quinbaya National Park, La Suiza, 1900 m, 13.III.2011, leg. J. OLÁH Jr. (1 male, OPC). : Tinalandia Nature Reserve, West Andean Slope, 600 m, 150 km near Quito 15.X.2010, light, leg. J. OLÁH Jr. (1 male, OPC).

sp. n. (Figs 283–287)

Diagnosis – This species has very slender and arching phallic organ similarly to the lateral phallic profile of S. peruana (MARTYNOV, 1912), S. petasata FLINT, 1981 and S. sudara sp. n. The new species is distinguished by having a spine fork on the phallic apex.

Description – Male. Pale brown, forewing length 6 mm. Maxillary palp formula 1-2-4-3-5. Tibial spurs 142. Cuticular structure of sternum V glands small with long anterolateral filament. Forewing membrane patterned: apical fourth darker and zigzag bordered; nygmal and thyridial point as well as chord (anastomosis) of crossveins s, r-m, m, m-cu and cu dark. 284

283

286 285

287

Smicridea (Rhyacophylax) villa sp. n., holotype, male genitalia: 283 = lateral view; 284 = left gonopod, ventral; 285 = dorsal; 286 = phallus lateral; 287 = phallus apex, ventral

Male genitalia. Segment IX with high staked ventrum in lateral view, anterodorsum indistinct; dorsal keel rounded covered with microtrichiae; intersegmental profile between segments IX and X sharp angled; ventromesally with short posterior triangular lobe. Seg- ment X long, lateral lobes straight long tapering; cerci longitudinally ovoid, located middle;

Annls hist.-nat. Mus. natn. hung. 104, 2012 296 J. Oláh & K. A. Johanson vestigial paraproct forming heavily sclerotised pair of ventrolateral straps on segment X directed downward from apical margin and fused to suture on ventrum IX, functioning as sclerotised phallic guides encircling above phallic organ. Basal segment of gonopods, coxopodite high with low basal fourth in lateral view; harpagones long slender parallel-sided, flat sagittaly and slightly bending mesad. Phallotheca with vertical inflated basal portion turning gradually to stem, and arching together; phallotheca constricted midway; apex with a pair of straight spines forming a forklike very distinct structure; apex with asperites of dorsolateral patch and ventromedian serrulate row of spicules; internal sclerotised portion of ejaculatory duct S-forming in lateral view bearing a pair of small internal spines.

Material examined – Holotype, male: : Chontachaca, Kosnipata-Cusco, 13°01’ 25”S, 71°28’03”W, 700 m, humid subtropical forest, Malaise trap 13–30.VII.2005, leg. J. FAVIOLA MONTES CARLOS (NRM). Paratypes: same data as holotype (1 male, OPC). San Martin Prov., creek crossing rd. Tarapoto-Yurimaguas, ca. 30 km (rd.) NE Tarapoto, 6° 24.904’S, 76°18.756’W, light, loc. 08, 8.I.2009, leg. K. A. JOHANSON &T.MALM (10 males, NRM, 7 males, OPC). San Martin Prov., La Catarata de Ahuashiyascu, 6°27.544’S, 76°18.192’ W, light, loc. 07, 7.I.2009, leg. K. A. JOHANSON &T.MALM (15 males, NRM, 11 males, OPC). San Martin Prov., creek crossing rd. Juan Guerra-Chazuta, 14 km (rd.) E Colombia Bridge, 6°35.594’S, 76°13.172’W, light, loc. 09, 9.I.2009, leg. K. A. JOHANSON &T.MALM (2 males, NRM).

Etymology – Villa, from “villa”, fork in Hungarian, refers to the backward directed pair of straight spines on the phallic apex.

REFERENCES

EVENHUIS N. L. 2007: Helping solve the “other” taxonomic impediment: completing the eight steps to total enlightenment and taxonomic Nirvana. – Zootaxa : 3–12. FLINT O. S. Jr. 1974: Studies of Neotropical caddisflies, XVII: The genus Smicridea from North and Central America (Trichoptera: Hydropsychidae). – Smithsonian Contri- butions to Zoology : 1–65. FLINT O. S. Jr., HOLZENTHAL R. W. & HARRIS S. C. 1999: Catalog of the Neotropical Caddisflies (Trichoptera). – Ohio Biological Survey, Columbus, Ohio, 239 pp. FLINT O. S. Jr., MCALPINE J. F. & ROSS H. H. 1987: A revision of the genus Leptonema Guérin (Trichoptera: Hydropsychidae: Macronematinae). – Smithsonian Contribu- tions to Zoology : 1–193. HOLZENTHAL R. W., BLAHNIK R. J., PRATHER A. L. & KJER K. M. 2007: Order Trichoptera Kirby, 1813 (Insecta), caddisflies. – Zootaxa : 639–698. MARTIN R. P. A. & J. V. SGANGA 2011: Smicridea McLachlan (Trichoptera: Hydropsy- chidae) from northwestern Argentina and Bolivia: new species, redescription, association and new records. – Journal of Natural History (35–36): 2219–2234. MARTYNOV A. V. 1912: On two collections of Trichoptera from Peru. – Annuaire du Musée Zoologique de l’Académie Impériale des Sciences de Saint Pétersbourg : 1–40.

Annls hist.-nat. Mus. natn. hung. 104, 2012 New Neotropical Macronematinae and Smicrideinae (Trichoptera: Hydropsychidae)297

MORSE J. C. (ed.) 2012: Trichoptera World Checklist. – http://entweb.clemson.edu/ database/trichopt/index.htm [Accessed 29 May 2012.] MOSELY M. E. 1933: A revision of the genus Leptonema (Trichoptera). – British Museum (Natural History), London, 69 pp. NIELSEN A. 1957: A comparative study of the genital segments and their appendages in male Trichoptera. – Det Kongelige Danske Videnskabernes Selskab, Biologiske Skrifter (5): 1–159. OLÁH J. & JOHANSON K. A. 2008: Reasoning an appendicular and functional caddisfly genital terminology. – Braueria : 29–40. SCHMID F. 1968: La famille des Arctopsychides (Trichoptera). – Mémoires de la Société Entomologique du Québec : 1–84. SCHMID F. 1970: Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera). – Mémoires de la Societé Entomologique du Canada : 1–230+LII plates.

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