﻿The Trichoptera of Panama XXIV. Fifteen new species and two new country records of the caddisfly genus Neotrichia (Trichoptera, Hydroptilidae), with a key to all known Panamanian species

﻿Abstract In this paper, 15 new species of microcaddisflies in the genus Neotrichia Morton, 1905 (Trichoptera, Hydroptilidae) from Panama are described and illustrated: Neotrichiaabrebotellasp. nov.; Neotrichiacandelasp. nov.; Neotrichiacodazasp. nov.; Neotrichiaemberasp. nov.; Neotrichiaflennikenisp. nov.; Neotrichiahondasp. nov.; Neotrichialandisaesp. nov.; Neotrichialenatisp. nov.; Neotrichiamindyaesp. nov.; Neotrichiapanamensissp. nov.; Neotrichiaparajarochitasp. nov.; Neotrichiaparaxicanasp. nov.; Neotrichiasnixaesp. nov.; Neotrichiaspanglerisp. nov.; Neotrichiaveraguasensissp. nov. In addition, two new country records are presented: Neotrichiaminutisimella (Chambers, 1873) and Neotrichiavibrans Ross, 1944. Finally, the male of N.vibrans is re-illustrated, the female is illustrated and descriptive information given, and a key is provided to the males of all current Neotrichia species in Panama. There are now 45 species of Neotrichia and a total of 525 Trichoptera species recorded from Panama.


Introduction
A concentrated effort during the last eight years (2015)(2016)(2017)(2018)(2019)(2020)(2021)(2022)(2023) has almost doubled the known caddisfly fauna of Panama from 257 to 508 species distributed among 15 families and 56 genera (e.g., Armitage et al. 2015;Harris and Armitage 2019;Harris et al. 2023).Concomitant with the increase in species was the additional gain of two families and 11 genera (e.g., Armitage et al. 2016;Armitage and Harris 2018;Armitage et al. 2022).These increases were made possible by adoption of an integrated sampling scheme involving multiple methods (primarily UV light traps and Malaise traps in combination) employed monthly for extended periods (usually January through June) at each collection site.Our knowledge of the family Hydroptilidae has been a particular beneficiary of this approach, generating more new species to science and new country records than for all other caddisfly families combined.No microcaddisfly taxa have supported this increase more than the genus Neotrichia Morton, 1905. ZooKeys 1188: 47-90 (2024), DOI: 10.3897/zookeys.1188.111346 Steven C. Harris et al.: New species and records of Neotrichia from Panama Before 2015, only three species of Neotrichia were known from Panama.Since then, we have added 25 new species and new country records (Table 1).Collections from selected national parks and protected areas, as well as public and private landholdings, were made during 2017-2023 and yielded 15 new species and two new country records of Neotrichia.Combined with the 28 known taxa, there are now 45 species of Neotrichia found in Panama, with 42 of them added since 2014 (31 new species and 11 new country records).The incremental additions to Panama's Neotrichia fauna are presented in Table 1, with associated literature references provided.We also re-illustrate the male of Neotrichia vibrans Ross, 1944 and provide descriptive text and illustrations for the female of that species.Finally, we provide a key to males of all 45 species of Neotrichia known from Panama.
The Aquatic Invertebrate Research Group (AIRG) at the Universidad Autónoma de Chiriquí (UNACHI) and its Museo de Peces de Agua Dulce e Invertebrados (MUPADI) is currently focused on increasing our knowledge of Trichoptera (caddisflies) and Plecoptera (stoneflies) in Panama.Toward that goal, it has secured registered projects for these two orders of aquatic insects.The new taxonomic information presented in this paper are a direct result of executing these projects.* Prior to 2015, three species of Neotrichia were known from Panama: N. flowersi Harris, 1990;N. malickyi Harris (in Harris and Tiemann 1993); and N. tauricornis Malicky, 1980.

Protected areas or private landholdings
Some of the collection sites referenced in this paper are found on private landholdings or national protected areas.These include the following study areas.
They are referred to in a number of ways throughout the text.We are constrained from unifying the naming system for these because the collection label names and administrative codes for the various locations cannot be changed.Therefore, we present the Spanish names for each study area as a header followed by any abbreviations or label names used in parentheses.In the text that follows each study area, we provide the English version of the study area name, if in common usage, as additional information.
Parque Nacional Santa Fe

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72,636 ha.Occupying land on both the Caribbean and Pacific slopes, more than 95% of the parks area is covered with tree species which are evergreen, maintaining their leaves all year round.
Parque Nacional Altos de Campana All of the national parks and forest reserves indicated above are under the stewardship of the Ministerio de Ambiente de Panamá (MiAmbiente) and protected from logging and agriculture.The streams sampled under this project are 1 st to 3 rd order in size, are of good water quality, and are bordered by extensive, forested riparian corridors.The private landholdings above include some agricultural activities; however, all or almost all streams are associated with well-vegetated riparian corridors as they traverse the properties.Most of the streams are found in major watersheds (cuencas), including cuencas 102, 108, 115, 117, 132, 134, and 138, which are characterized in Cornejo et al. (2017).Cuenca 097 (Río Calovébora watershed; Caribbean drainage for Santa Fe National Park), Cuenca 104 (Río Escárrea; Pacific drainage), and Cuenca 116 (rivers between the Tabasará and the San Pablo; Pacific drainage), however, are not included in that book.

Methods
Collections from selected national protected areas, private landholdings, and public areas were made during 2017-2023.Both Malaise and UV light traps were used for collecting aquatic insects from streams in the national parks and protected areas of Panama.Single, overnight collections were made using UV light traps (Calor and Mariano 2012).Generally, multiple-night collections were made employing Malaise traps over four-day periods.Specimens were prepared and examined following standard methods outlined in Blahnik and Holzenthal (2004).Male genitalia were soaked in 5% KOH overnight, and washed in weakly acidified alcohol prior to examination under a dissecting scope.For illustrations, specimens were slide-mounted and viewed at 250× magnification.
Morphological terminology used for male genitalia generally follows that of Marshall (1979) and classification within the Hydroptilidae follows Thomson (2023).Paired structures are described in the singular for simplicity.Although technically, segments V through X are not part of the genitalia, traditionally descriptions of segments VII through X have been included under the genitalia heading.We follow that practice here.If segments V and VI have distinct features, they are discussed under the male description.Total length of specimens provided in descriptions represents the length from the tip of the head to the tip of the forewing.Altitude values are given in meters above sea level (m a.s.l.).Maps were created using QGIS software, version  Holotypes listed in this publication are deposited in the University of Panama's Museo de Invertebrados (MIUP) or MUPADI.Paratypes and other specimens are deposited in MUPADI, the University of Minnesota Insect Collection (UMSP), or the first author's reference collection (SCH).The species listed below are in alphabetic order.

Taxonomy-new species
Genus Neotrichia Morton, 1905 Remark.The genus Neotrichia (Hydroptilidae, Neotrichiinae) has a New World distribution with 211 species known from North, Central, and South America and the West Indies (Armitage et al. 2022;Gama-Neto and Passos 2023;Harris et al. 2023;Thomson 2023).In the Neotropics, there are 174 species recorded (Armitage et al. 2022;Gama-Neto and Passos 2023;Harris et al. 2023;Thomson 2023).Herein we describe and illustrate 15 new species and report two new country records for Panama.Diagnosis.Neotrichia abrebotella sp.nov.appears to be a member of the N. caxima group of Keth et al. (2015) based on the short inferior appendage with resemblance to N. atopa Thomson &Armitage, 2018 andN. caxima Mosely, 1937.It differs from N. caxima in the lack of sclerotized phallic processes and from N. atopa in the bifid appearance of the inferior appendage in lateral view and the narrow bracteole.
Description.Male.Total length 1.7-1.9mm (n = 3), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 2).Abdominal segment VIII annular.Segment IX in lateral view rounded posteriorly, narrowing dorsally, anteriorly narrowing to elongate mesal apodeme; in dorsal and ventral views quadrate, deeply incised anteriorly.Tergum X short, truncate posteriorly, tapering on lateral margins; in lateral view truncate posteriorly.Subgenital plate in lateral view wide subapically, narrowing distally to rugose, downward projecting process; in ventral view wide, truncate posteriorly with median lobe and pair of stout setae, rounded on lateral margins.Bracteole narrow basally, slightly widening distally; in dorsal and ventral views lobate.Inferior appendage short and narrow, bifid posteriorly, lower portion longer than upper; in ventral view square basally, distally bifid, dorsalmost process narrowing mesally to acute apex, ventralmost process tubular.Phallus tubular, wide basally, constricted at midlength and bearing thin paramere encircling shaft, apically thin over length.
Etymology.The species name abrebotella (bottle-opener) derives from Spanish, referring to the distinctive appearance of the inferior appendage.The name is a noun in the nominative singular standing in apposition.Diagnosis.Neotrichia candela sp.nov. is a member of the N. canixa group of Keth et al. (2015) based on the posterior horns from tergum X, forked bracteoles, and the bifid inferior appendage.The new species appears to be similar to N. bika Oláh & Johanson, 2011 from French Guiana on the basis of the structure of the bracteoles and the horns of the tenth tergum.The new species is separated by the structure of the phallic apex, which has more elongate forking than that of N. bika, and by the lateral appearance of the inferior appendage and subgenital plate.
Description.Male.Total length 1.7 mm (n = 1), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 3).Abdominal segment VIII annular.Segment IX in lateral view generally ovate, rounded anteriorly and posteriorly, fused with segment X dorsally with seta-bearing lobe posterolaterally; in ventral view anterior margin deeply incised, posterior margin with thin, elongate mesal extension; in dorsal view incomplete posteriorly.Tergum X basally fused with segment IX, wide basally, tapering posteriorly forming pair of stout, symmetrical horns; in lateral view segment X elongate, dilated distally with distal horn short and tapering to acute apex.Subgenital plate in lateral view, wide basally, tapering distally to bifid apex, upper arm short, with acute apex, lower portion elongate and crescent-shaped; in ventral view triangular-shaped, apex with pair of thin lateral lobes bearing stout setae, mesally with short distal triangle and ventral sclerotized process.Bracteole in lateral view wide anteriorly, bifid posteriorly, dorsal branch twice as long as ventral branch, each bearing terminal seta; in ventral and dorsal views both branches wide basally, tapering to rounded apices.Inferior appendage rectangular in lateral view, subapically with dorsal sclerotized points, basal process elongate and thin; in ventral view bifid, outer process slightly wider basally, curving and tapering to rounded apices, inner process fused basally and narrow over length which is ~ 1/2 of outer process, bearing elongate seta apically.Phallus tubular, constricted at midlength and bearing thin paramere encircling shaft, apex forked with elongate processes, lower of which sharply curves laterally, ejaculatory duct protruding at base.
Etymology.This new species is named for the Río Candela in western Chiriquí Province where the species was collected.The name is a noun in the genitive case.from Peru, based on the sharing of the distinctive pair of dark spines at the phallic apex.The new species is separated by the elongate, angled inferior appendage, and the lack of a mesal process from the posterior margin of segment IX. Description.Male.Total length 1.3-1.5 mm (n = 12), 17 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 4).Abdominal segment VIII annular.Segment IX in lateral view with shallow dorsal and ventral emarginations posteriorly, anteriorly tapering to acute apex; dorsally and ventrally deeply incised anteriorly, shallow emarginations posteriorly.Segment X short and lobate in lateral view; in dorsal view apically truncate, flared laterally.Subgenital plate in lateral view, wide basally, abruptly tapering apically on venter forming acute spine, basal portion developed laterad, widening distally and flaplike; in ventral view, quadrate with apicomesal extension bearing stout setae laterad, lobate laterally.Bracteole in lateral view wide basally, tapering distally to rounded apex; in ventral and dorsal views short, tapering to rounded apex.Inferior appendage wide basally with long dorsal arm, narrowing and angled dorsad at midlength and bearing stout dorsal seta, widening distally to rounded apex; in ventral view wide basally, tapering distally to rounded apex, elongate dorsal arm wide at base, tapering distally to round apex.Phallus in dorsal view wide basally and apically, bearing short paramere encircling shaft at midlength, apex with pair of short dark spines; in lateral view apical spines are close together and slightly curving dorsally.
Etymology.The species name codaza (to elbow) derives from Spanish, referring to the bent inferior appendage in lateral view.The name is a noun in the nominative singular standing in apposition.Diagnosis.This new species, with a pair of apical phallic spines, would appear to be another member of the N. collata group of Keth et al. (2015), with similarity to N. anahua (Mosely, 1937) from Mexico, and N. kurtika Oláh & Johanson, 2011 from French Guiana.Unlike these species, N. embera sp.nov.Harris, Armitage & Ríos has a posterolateral process from segment IX and bifid inferior appendages.

Neotrichia embera
Description.Male.Total length 1.2 mm (n = 1), 17 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 5).Abdominal segment VIII annular.Segment IX in lateral view anteriorly tapering to short upturned apodeme, posteriorly with medial lobe, distally rounded with posterolateral triangular process; ventrally with anterior and posterior margins incised mesally, posteriorly with thin lateral processes; dorsally truncate posteriorly.Segment X elongate, triangular in lateral view; in dorsal view tapering distally to rounded apex.Subgenital plate in lateral view narrow basally, ventral margin sinuate with lobes and points, apex with stout seta, upper margin straight; in ventral view rectangular, widening posteriorly with stout setae on distal margins, rounded laterally.Bracteole in lateral view narrow basally, widening distally to rounded apex; in ventral and dorsal views straight, narrow over length.Inferior appendage bifid, ventral arm narrowly triangular, dorsal arm ovate and same length as lower arm; in ventral view divergent and separated, lateral arms narrow over length, inner arms slightly triangular, rounded apically.Phallus in dorsal view tubular, wide basally and distally, bearing pair of stout spines apically, short paramere encircling shaft at midlength.
Etymology.This species is named for the indigenous Embera people of Darién Province, where the species was collected.The name is a noun in the genitive case.Diagnosis.Neotrichia flennikeni sp.nov.appears to be a member of the N. vibrans group of Keth et al. (2015) based on the posterolateral process of segment IX, and the tapered inferior appendage.The new species appears to be similar to N. angulata Flint, 1983 from Uruguay which also has an elongate, tapering inferior appendage and a phallus bearing a long, stout spine.The new species is distinguished by the knob-like posterolateral processes from segment IX and by the tapering, rather than angled, appearance of the inferior appendage.Description.Male.Total length 1.8 mm (n = 1), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 6).Abdominal segment VIII annular.Segment IX in lateral view narrow, tapering anteriorly, posteriorly with thin posterolateral process; ventrally with deep rounded posterior excision; dorsally with narrow posterior incision, posterolateral processes sclerotized and knobbed apically.Segment X narrowing posteriorly to thin shelf; in dorsal view wide basally, tapering distally to truncate apex.Subgenital plate in lateral view, narrow basally, widening distally, with acute, posteriorly projected dorsal spine, sclerotized knob posteroventrally; in ventral view narrowing basally, with pair of apical lobes each bearing stout seta.Bracteole in lateral view widening posteriorly to rounded apex; in ventral and dorsal views narrow over length.Inferior appendage in lateral view elongate and tapering, curving upward at midlength; in ventral view wide basally, sharply angled at midlength, then narrowing to acute apex which projects inward.Phallus tubular, constricted at midlength and bearing thin paramere encircling shaft, apex rectangular with elongate, stout medial spine.
Etymology.This species is named after Donald G. Flenniken of eastern Ohio, a teacher, naturalist, and part-time milkman who taught the second author (at age of 11) how to identify mammals from their skull bones/dentition and how to use taxonomic keys-a brief encounter that has led to a lifetime of taxonomic pursuits and pleasures.The name is a noun in the genitive case.Diagnosis.Neotrichia honda sp.nov.does not fit well in any of the species groupings established by Keth et al. (2015).The new species has forked sclerites at the apex of the phallus and has an overall resemblance to N. gilaensis Keth et al., 2015 from Arizona which is in the N. collata group, but it is also similar to N. yavesia Bueno-Soria, 2010 from Mexico in the N. vibrans group, based on the short, rectangular inferior appendages.The new species is separated by the elongate, forked sclerites of the phallus apex, the elongate anterior apodeme of segment IX, and by the sclerotized posterolateral processes from segment IX.
Description.Male.Total length 1.5 mm (n = 1), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 7).Abdominal segment VIII annular.Segment IX in lateral view anteriorly tapering to elongate apodeme, posteriorly rounded with posterolateral process, which is wide basally, tapering distally to an acute downward turning hook; dorsally and ventrally deeply incised anteriorly, ventrally with deep lateral incisions posteriorly, creating an elongate mesal, triangular lobe; dorsally with posterolateral processes narrow over length, curving slightly inward apically.Segment X elongate and thin in lateral view; in dorsal view wide basally and fused with segment IX, tapering to rounded apex.Subgenital plate in lateral view, wide basally, gradually tapering apically to rounded apex bearing elongate seta; in ventral view narrowly triangular, pair of stout setae apically.Bracteole in lateral view wide basally, then narrowing distally to truncate apex; in ventral and dorsal views narrow over length and curving mesally.Inferior appendage sclerotized and rectangular, wide basally, narrowing at midlength, then abruptly tapering to rounded apex; in ventral view triangular, rounded apically, basally with finger-like lobe.Phallus in dorsal view wide basally, bearing short paramere encircling shaft at midlength, apex divided into pair of elongate, sclerotized rods, which are nearly equal in length.Diagnosis.Neotrichia landisae sp.nov.belongs to a cluster of similar Panamanian species with characteristic elongate posterolateral processes from abdominal segment IX, including N. tatianae Armitage &Harris, 2018 andN. yayas Armitage &Harris, 2020.The new species is separated from these species by the linear posterolateral processes from segment IX, the truncate inferior appendage, and the narrow subgenital plate in ventral view.
Description.Male.Total length 1.5-1.7 mm (n = 15), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 8).Abdominal segment VIII annular.Segment IX in lateral view anteriorly rounded, wide posteroventrally, narrowing dorsally, pair of elongate, thin posterolateral processes, which narrow distally; in dorsal view fused posteriorly with segment X, pair of small setaceous lobes laterally, posterolateral processes sclerotized and elongate, but dissimilar in length, anteriorly with narrow mesal incision; in ventral view with deep lateral incisions.Tergum X basally fused with segment IX, posteriorly elongate, tapering to rounded mesal lobe; in lateral view wide basally, tapering distally to acute apex.Subgenital plate in lateral view wide basally, tapering distally to narrow shelf bearing stout seta; in ventral view narrow over length, pair of setae apically.Bracteole absent or represented by thin, elongate seta-bearing process on dorsal margin of segment IX. Inferior appendage short, wide basally, widening subapically, spinose and rugose on distal margin, thin ventral process basally; in ventral view elongate and narrow, apical point turning inward, spinose subapically on mesal margin, base of inferior appendage and processes from segment IX superimposed, short triangular mesal process bearing stout seta apically.Phallus tubular, constricted at midlength and bearing thin paramere encircling shaft, apically rounded with ejaculatory duct protruding.
Etymology.This species is named for Senora Marietta Isabel Landis, owner of Reserva Privada Landis, who managed all collection events.The name is a noun in the genitive case.

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species is separated by the presence of a basal process from the inferior appendage, narrow bracteole, and the ventral location and shape of the posterolateral process from segment IX. Description.Male.Total length 1.4 mm (n = 1), 18 antennal segment, wings and body brown in alcohol.Genitalia (Fig. 9).Abdominal segment VIII annular.Segment IX in lateral view narrow, tapering anteriorly to mesal apodeme, truncate posteriorly, with thin process posterolaterally, tapering ventrad and incised on posterior margin; in ventral view narrow, sinuate posteriorly; in dorsal view deeply incised mesally, forming rounded lateral lobes.Segment X seemingly fused with segment IX in lateral view, tapering downward to posterior projecting shelf with pointed tip; in dorsal view short, rounded distally, with truncate apex and sclerotized margins.Subgenital plate triangular in lateral view; in ventral view quadrate, lateral apices indented with stout seta, small mesal downturned lobe.Bracteole narrow in lateral view, slightly widening distally; in ventral and dorsal views narrow over length.Inferior appendage sclerotized, tapering distally and slightly curving upward at midlength, dorsobasally with thin process tipped with stout seta; in ventral view tapering distally to rounded apex, knoblike basal process on inner margin.Phallus tubular, constricted near midlength and bearing thin paramere encircling shaft, posteriorly thin and rectangular, ejaculatory duct lightly sclerotized.
Etymology.This species is named in memory of David R. Lenat of Raleigh, North Carolina, our friend and colleague, who was an avid researcher in taxonomy and aquatic bioassessment, and who made numerous important contributions to our knowledge of chironomids, stoneflies, caddisflies, mayflies, and many other groups of aquatic insects.The name is a noun in the genitive case.Diagnosis.Neotrichia mindyae sp.nov. is another member of the N. canixa group of Keth et al. (2015) based on the posterior horns from tergum X, forked bracteoles, and the bifid inferior appendage.The new species appears to be most similar to N. maya Harris & Flint, 2016 from Belize on the basis of the structure of the bracteole and the horns of tergum X.The new species is separated by the structure of the phallic apex, which while forked, has the one process extremely long, similar to that of N. maria Bueno-Soria & Hamilton, 1986 from Mexico, and by the unequal forking of the bracteole, which are uniformly forked in N. maria.
Segment IX in lateral view generally ovate, rounded anteriorly and posteriorly, fused with segment X dorsally, and bearing lateral setal lobes posteriorly; in ventral view anterior margin deeply incised, posterior margin with triangular mesal extension; in dorsal view with pair of posterolateral lobes.Tergum X basally fused with segment IX, posteriorly produced into pair of thick, symmetrical horns which nearly touch mesally; in lateral view distal horns are thin and tapering to acute apices.Subgenital plate in lateral view, wide basally, tapering distally to downturned apical hook; in ventral view wide basally, rounded laterally to apicomesal extension.Bracteole in lateral view wide anteriorly, bifid posteriorly, dorsal branch elongate with apical seta, ventral branch greatly reduced with apical seta; in ventral and dorsal views both branches wide basally, tapering to rounded apices.Inferior appendage bifid laterally, each thin arm wide basally and tapering distally to rounded apices bearing long seta; in ventral view bifid, outer process wide basally, curving and tapering to acute apices, inner process fused basally and narrow over length which is equal to outer process bearing elongate seta apically.Phallus tubular, wide basally, constricted at midlength and bearing thin paramere encircling shaft, apex with elongate lateral spike, smaller spike near base, ejaculatory duct not protruding.Distribution.Panama: Darién Province (Darién National Park).
Etymology.This species is named for the sister of the first author, Melinda "Mindy" Harris Haupt, who used to help her brother collect bugs when they were both much younger.The name is a noun in the genitive case.Diagnosis.Neotrichia panamensis sp.nov. is another member of the N. canixa group of Keth et al. (2015) based on the posterior horns from tergum X, forked bracteole, and the bifid inferior appendage.The new species appears to be similar to N. alsa Oláh & Johanson, 2011 from Peru on the basis of the bracteoles and phallic apex, but the short inferior appendages are more like those of N. tauricornis Malicky, 1980 which occurs throughout the Caribbean islands, Panama, and Colombia, but the subgenital plate and bracteoles are much different in the new species compared to that of N. tauricornis.

Neotrichia panamensis
Description.Male.Total length 1.5-1.7 mm (n = 14), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 11).Abdominal segment VIII annular.Segment IX in lateral view ovate, rounded anteriorly and sinuate posteriorly, fused with segment X dorsally, bearing a setae-bearing lobe dorsally; in ventral view anterior margin deeply incised, posterior margin sinuate, triangular mesal extension.Tergum X basally fused with segment IX, rectangular basally, pair of thin, symmetrical horns distally; in lateral view segment X is lobate, with distal horn saber-like.Subgenital plate in lateral view, wide basally, truncate distally with ventral hook tapering to acute apex; in ventral view wide basally, slightly curving to rounded apex, with mesal process flanked by stout setae.Bracteole in lateral view wide anteriorly, slightly bifid posteriorly, dorsal branch elongate and tipped with seta, ventral branch vestigial, represented by pair of sharp points; in ventral and dorsal views lower branch represented by short knob, tapering distally.Inferior appendage short and bifid, wide basally tapering distally, basal process triangular; in ventral view bifid, outer process subrectangular, apex with mesal points, inner process same length as outer, fused and wide at base, tapering distally bearing elongate seta apically.Phallus tubular, constricted at midlength and bearing thin paramere encircling shaft, apex divided into three processes, lower process small, distally processes elongate, ejaculatory duct not protruding at base.Etymology.This species is named for the Republic of Panama where the species was collected.The name is a noun in the genitive case.Diagnosis.Neotrichia parajarochita sp.nov. is another member of the N. canixa group of Keth et al. (2015) based on the posterior horns from tergum X, forked bracteole, and the forked apex of the phallus.In many respects, the new species appears to be similar to N. jarochita Bueno-Soria, 1999 andN. palitla Harris &Flint, 2016, both of which occur in Mexico, on the basis of the asymmetrical horns of tergum X and the structure of the bracteole.Unlike these two species, N. parajarochita has a sclerotized posterolateral process from segment IX and the inferior appendages are divergent in ventral view.
Description.Male.Total length 1.4 mm (n = 1), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 12).Abdominal segment VIII annular.Segment IX in lateral view ovate, rounded anteriorly and sinuate posteriorly, setae-bearing lobe dorsally and fused with segment X, with upturned posterolateral process mesally, narrowing distally to serrate apex; in ventral view anterior margin emarginate, posterior margin incised mesally, with an oval mesal extension, and posterolateral processes laterad, narrow over length and curving.Tergum X basally fused with segment IX, quadrate, posterior horns asymmetrical, left horn twice as long as right; in lateral view segment X is truncated, distal horn elongate and tapering to acute apex.Subgenital plate in lateral view, wide basally, tapering distally to acute apex, which is slightly downturned; in ventral view roundly triangular, abruptly narrowed distally forming rectangular mesal extension flanked by pair of setae.Bracteole in lateral view wide anteriorly, bifid posteriorly, thin dorsal branch slightly longer than triangular ventral branch, each tipped with seta; in ventral and dorsal views wide basally, tapering distally.Inferior appendage short in lateral view, widening at midlength, then tapering to rounded apex; in ventral view bifid, outer process wide basally, narrowing distally and strongly diverging, rectangular apically with lateral spike, inner process shorter and narrower than outer process.Phallus tubular, constricted at midlength and bearing thin paramere encircling shaft, apex forked, lower fork strongly curving, ejaculatory duct protruding subapically.
Etymology.This species is named for its resemblance to Neotrichia jarochita.The name is an adjective used as a substantive in the genitive case.species is most similar to N. xicana (Mosely, 1937), reported from Mexico and Panama (Holzenthal and Calor 2017), based on both having a posterolateral process from segment IX, which is lobate apically.However, in the new species this process is very short, and the phallic apex is multi-forked.Description.Male.Total length 1.6-1.8mm (n = 3), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 13).Abdominal segment VIII annular.Segment IX in lateral view generally ovate, rounded anteriorly, setae-bearing lobe dorsally, posteriorly fused with segment X dorsally, incised posteroventrally, short posterolateral process with multiple small lobes apically; in ventral view posterior margin deeply incised laterally, anteriorly margin broadly incised; in dorsal view posterolateral process from IX visible as a curving, tapering process with subapical lobes laterally.Tergum X basally fused with segment IX, posteriorly produced into pair of thick, symmetrical horns; in lateral view distal horn is thin and tapering to acute apex.Subgenital plate in lateral view, wide basally, tapering apically to rounded apex, with acute downward pointing spike; in ventral view wide basally, rounded laterally to apicomesal extension bearing stout setae laterad.Bracteole in lateral view wide anteriorly, bifid posteriorly, dorsal branch ~ 2× as long as ventral branch; in ventral and dorsal views both branches wide basally, tapering to rounded apices.Inferior appendage wide basally with narrow basal process ventrally, abruptly narrowing at midlength and curving upward to thin, truncated apex; in ventral view bifid, outer process elongate, wide basally and tapering to acute apices with inner spike, inner process fused basally and narrow over length, shorter than outer process.Phallus tubular, constricted at midlength and bearing thin paramere encircling shaft, apex divided into four spine-like processes, ejaculatory duct protruding subapically.
Etymology.This species is named for its overall resemblance to Neotrichia xicana.The name is an adjective used as a substantive in the genitive case.
Diagnosis.Neotrichia snixae sp.nov. is a member of the N. canixa group of Keth et al. (2015) based on the posterior horns from tergum X and the forked bracteole.The horns are asymmetrical, which is similar to that seen in N. malickyi Harris & Tiemann, 1993 from Panama and N. jarochita Bueno-Soria, 1999 from Mexico, both of which have the right horn longer than that of the left.The new species differs in the presence of a sclerotized posterolateral process from segment IX, which is also seen in N. juani Harris &Tiemann, 1993 from Texas andN. unamas Botosaneanu (in Botosaneanu andAlkins-Koo 1993), which occurs in Panama, and in the shape of the end of the phallus.
Description.Male.Total length 1.9 mm (n = 2), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 14).Abdominal segment VIII annular.Segment IX in lateral view anteriorly rounded, incomplete posteriorly, fused with X dorsally, mesally with sclerotized posterolateral process which narrows distally; in dorsal view fused posteriorly with X, pair of setaceous lobes laterally, anteriorly with rounded incision, lateral processes dissimilar in length and   process; ventrally deeply incised anteriorly, posteriorly with lateral incisions forming mesal ovate structure; posterolateral processes thin, tapering to acute apices; dorsally with posterior shallowly incised laterally, setose mesally.Segment X in lateral view tapering to acute distal point; in dorsal view wide basally, rounded laterally to truncate apex.Subgenital plate in lateral view narrow, apex downturned and acute; in dorsal and ventral view wide basally, apex with mesal emargination, flanked by pair of setae.Bracteole in lateral view narrow basally, widening distally to rounded apex; in ventral and dorsal views nearly parallel-sided, curving on outer margin.Inferior appendage short, wide basally, tapering distally to broadly pointed apex; in ventral view rectangular, apex tapered to rounded point.Phallus in dorsal view tubular, bearing short paramere encircling shaft at midlength, apex with pair of elongate sclerotized rods fused basally, one rod ~ 1/2 length of other and curved.
Etymology.The species name honors the memory of Dr Paul Spangler of the National Museum of Natural History, who collected some of the specimens.The name is a noun in the genitive case.Other material examined.Panama • ♂; ibid., except Río Piedra de Moler; PSP-SCB-PNSF- C097-2017-012;8.56553°N, 81.18817°W;340 m a.s.l.;20.iv.2017;MIUP (in alcohol).

Neotrichia veraguasensis
Diagnosis.This new species, which lacks spines at the phallic apex, is placed in the N. okopa group of Keth et al. (2015), with closest similarity to N. abbreviata Flint, 1983 from Uruguay and N. okopa Ross, 1939 from throughout North America.The new species is separated by the triangular inferior appendage in lateral view, and the lack of a posterolateral process from segment IX.
Description.Male.Total length 1.6 mm (n = 2), 18 antennal segments, wings and body brown in alcohol.Genitalia (Fig. 16).Abdominal segment VIII annular.Segment IX in lateral view anteriorly tapering to short apodeme, posteriorly rounded, short truncate lobe posteroventrally; ventrally divided into two posterior bands; dorsally with anterior margin incised, posterior shallowly emarginate.Segment X triangular in lateral view; in dorsal view fused basally with segment IX, posteriorly sinuate on lateral margin and tapering to rounded apex.Subgenital plate in lateral view narrow basally, ventral margin with large sclerotized lobe at midlength, upper margin straight distally, abruptly tapered posteriorly to rounded apex bearing stout seta; in ventral view wide and rounded laterally to truncate apex, with setae laterally.Bracteole in lateral view parallel-sided to rounded apex; in ventral and dorsal views straight, narrow over length.Inferior appendage triangular and sclerotized; in ventral view divergent, lateral margin curving, inner margin straight, round lobe at midlength, basally with elaborate thin sclerites.Phallus in dorsal view tubular, bearing short paramere encircling shaft at midlength, apex rectangular with ejaculatory duct protruding mesally.Etymology.This species is named for Veraguas Province where the male holotype was collected.The name is a noun in the genitive case.Female (Fig. 18).The female of N. vibrans is slightly larger than the males, with total length from 2.0 to 2.5 mm compared to 1.8 to 2.0 mm, both having 18 antennal segments and the wings and body are brown in alcohol.Abdominal sternite VIII has a heavily sclerotized square plate (Fig. 18A), with a thin ridge on the posterior margin that narrows to an acute posteromesal point, which projects downward in lateral view, anteriorly the plate margin is rounded with a thin sinuate internal structure.The bursa copulatrix lies under the sclerotized plate of segment VIII and details are difficult to discern, but it is generally rectangular in shape, the genital chamber narrowing posteriorly to an acute point, genital chamber broadly incised anteriorly producing a pair of "feet" with heavily sclerotized inner margins, these "feet" extend posteriorly forming a mesal sclerite (Fig. 18B).

New country records
Note.This is a new country record for Panama.In addition, it is a significant southern extension of the species range, which formerly was northern Mexico.The male of this species is here re-illustrated (Fig. 17) because the figures in Keth et al. (2015) do not clearly depict the lateral view of this species.The posterolateral processes from abdominal segments IX and X are difficult to see in those drawings and the bracteole is obscured by the process from IX.As well, the figure for N. vibrans in Ross (1944) lacks a lateral view and the phallus is shown as the apex having a pair of lobes bearing stout spines.These lobes and spines, which are also used as characters in the diagnostic key in Ross (1944), belong to the apex of the subgenital plate, with the phallus lacking apical spines.
Females of other species from the N. vibrans group of Keth et al. (2015) also have a sclerotized plate on the sternum of segment VIII, including Neotrichia ♀ sp.B (in Botosaneanu and Alkins-Koo 1993), which was tentatively identified as N. armata Botosaneanu, 1993, N. iridescens Flint, 1964, andN. soleaferrea   the sternum of sternite VIII lacks a sclerotized plate.Also, with the exception of N. soleaferrea, the structure of the bursa copulatrix appears to be much different than that seen in N. vibrans.
The ventral plate on sternite VIII is not a characteristic unique to the N. vibrans group.Females in other groups have a distinctive sclerotized plate, e.g., N. margararitena Botosaneanu (in Botosaneanu and Viloria 2002) of the N. biuncifera group (Marshall 1979).Also, females for several members of the N. caxima group, including N. nesiotes Flint & Sykora, 1993, N. mentonensis Frazer & Harris, 1991, N. rasmusseni Harris & Keth, 2002, and N. armitagei Harris, 1991 share this trait.In addition, many members of the N. caxima group also have females with the structure of the bursa copulatrix very similar to that seen in the female of N. vibrans (Harris and Rasmussen 2010).
Considering the above variation and overlap of characters, more associations are needed to facilitate a detailed comparison and delineation of Neotrichia females before we can adequately diagnose any females definitively.The descriptive text for N. vibrans above is a contribution toward that future comparison.Finally, we were fortunate in being able to associate the female of N. vibrans with the male because individuals of both sexes were found, in quantity, together in the absence of any other congeners in a non-natural botanical garden site.Most natural stream sites we have sampled in Panama have 2-7 species present in the same sample, making associations more difficult.

Key to the males of Neotrichia in Panama
Species of Neotrichia are difficult to identify given their tiny size, which varies from 1 to 3 mm, and the complex genitalia, on which identifications are dependent.The small size often necessitates that specimens are mounted on slides and observed at high magnification.Drawings are likewise prepared from these slide-mounted specimens.Two broad characters are often useful in making species determinations: small terminal horns located on the apex of tergum X and the presence of a posterolateral process from segment IX, which is often sclerotized.When the terminal horns are present, characters of the bracteoles, inferior appendages and phallus become important.When a posterolateral process from segment IX is present, then the appearance of these processes and their location and point of origin, the bracteoles, phallus, and inferior appendages all provide useful characters.For example, the posterolateral process can be mesal as in N. honda and N. parajarochita, dorsal as in N. landisae and N. paraxicana, and ventral as in N. lenati.In some instances, what appears to be a process from segment IX is actually a part of the subgenital plate, or another structure.For simplicity and ease of using this key, we have termed what appears to be a posterior extension of segment IX as a posterolateral process.

Discussion
With the publication of this paper, there are now 226 species of Neotrichia which occur in North, Central, and South America and the West Indies (Armitage et al. 2022;Gama-Neto and Passos 2023;Harris et al. 2023;Thomson 2023).Of these, 31 species are known from Mexico and 50 species from Central America, with Panama's 45 species comprising the majority of the latter.We suspect that many of Panama's 33 endemic species of Neotrichia would lose that status once the Trichoptera fauna in the other Central American countries becomes better known.Other than Belize, with five endemic (and total) species, no other Central American country hosts any endemics.Three species are recorded from Nicaragua, one from Costa Rica, and none are recorded from El Salvador, Guatemala, and Honduras (Thomson 2023).Given the close relationship between the faunas of Costa Rica and Panama, we expect that they share many species.In South America, Brazil is the dominant country for this genus with 48 recorded species (Gama-Neto and Passos 2023; Santos 2023), and has the majority of the endemic species.It shares only two species with Panama (N.tauricornis Malicky, 1980;N. parabullata Harris & Armitage, 2015), so that the faunas of these two countries, though currently similar in size, are more or less distinct.
Given the rapid increase in Neotrichia species we have found in Panama during the last eight years (42 species, with ~ 75% of them new to science), we are no longer surprised, but still delighted, at what is revealed in our samples.With most of Panama as yet unexplored, the possibility of 75, or even 100, species of this genus in Panama is not beyond the imagination.Whereas we are just now beginning to see second and third locations for some of the Neotrichia we have previously described, "new" species remain the central theme of our results.
It is well known that keys to species, while useful, are usually out of date as soon as they are written.However, we developed the key presented above out of self-defense to help manage the current list, to provide a framework for new keys in the future, and, most of all, to assist us in identifying and verifying new species as they appear.In Keth et al. (2015) an attempt was made to "further organize" primarily Nearctic species into species groups.However, some of the new Neotrichia species we are finding in Panama raise some doubts about the validity of those groupings.Increasing knowledge of the sampling universe visà-vis Neotrichia has not necessarily brought clarification in this regard, as might be expected.Other types of analyses, including molecular, need to be brought to bear to supplement what we think we know about this most interesting genus.

(
Santa Fe NP or PNSF; Fig. 1A: collection locations N and O) Located in the upper portion of the Santa Maria River basin in Veraguas Province, Santa Fe National Park lies near the Continental Divide and encompasses

(
Altos de Campana NP or PNAC; Fig. 1A: collection location U) Established in 1966, Altos de Campana National Park in Panama Oeste Province is the oldest park in Panama.Covering 1,950 ha, the park lies on the Pacific slope of Panama and is covered, in part, by humid tropical and premontane forests.Parque Nacional Darién (Darién NP or Darién National Park; Fig. 1A: collection location X) Darién National Park encompasses some 579,000 ha in Darién Province, adjacent to Colombia in southeastern Panama.Under formal protection since 1972, the Alto Darién Protection Forest became a national park in 1980, a World Heritage area since 1981, and, soon thereafter, a biosphere reserve by UNESCO.
Botosaneanu and Hyslop 1998).However, other members of this group have the females unidentified, or in the case of N. heleiosFlint, 1968
Type material.Holotype: ♂, Panama: Chiriquí Province: shape; in ventral view with deep mesal incision.Tergum X basally fused with segment IX, quadrate, posterior horns asymmetrical, right horn longer than left and thicker, curving inward distally; in lateral view ventral horn twice as long as dorsal horn.Subgenital plate in lateral view wide basally, tapering distally to divided apex, dorsally with stout seta, ventrally with elongate downturned process which is apically acute; in dorsal and ventral views wide basally, elongate posteromesally, rounded laterally.Bracteole in lateral view bifid posteriorly, ventral branch longer than dorsal, each tipped with stout seta; in dorsal and ventral views branches unequal in length, wide basally tapering distally.Inferior appendage wide basally, gradually tapering distally to rounded slightly upturned apex, narrow ventral process basally; in ventral view wide basally, curving on inner margin to rounded apex, mesal process tapering distally and bearing stout seta.Phallus tubular, constricted at midlength and bearing thin paramere encircling shaft, apically sclerotized with narrow apical incision forming unequal branches, similar in appearance to a can-opener; in lateral view apex divided into pair of elongate, thin processes.Named for Dr. Shannon Nix, friend and colleague of the first author in recognition of her outstanding teaching while at Clarion University and her many scientific contributions to ecology and mycology.The name is a noun in the genitive case.Aguirre leg.; UV light trap • ♂; ibid., except12.iii.2021• 2 ♂♂; ibid.,  except 12.iv.2021•11 ♂♂; ibid., except 6.x.2021 • 4 ♂♂; ibid., except 6.xi.2021 • 2 ♂♂; Darién Province; Cuenca 156, Chepigana District, PND, Río Tuira, Boca de Cupe; 8.01732°N, 77.72417°W; 150 m a.s.l.; 18.ii.1985;leg.not given; UV light trap (NMNH).