Research Article |
Corresponding author: Brian J. Armitage ( tobikera89@gmail.com ) Academic editor: Ana Previšić
© 2024 Steven C. Harris, Brian J. Armitage, Tomás A. Ríos González.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Harris SC, Armitage BJ, Ríos González TA (2024) 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. ZooKeys 1188: 47-90. https://doi.org/10.3897/zookeys.1188.111346
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In this paper, 15 new species of microcaddisflies in the genus Neotrichia Morton, 1905 (Trichoptera, Hydroptilidae) from Panama are described and illustrated: Neotrichia abrebotella sp. nov.; Neotrichia candela sp. nov.; Neotrichia codaza sp. nov.; Neotrichia embera sp. nov.; Neotrichia flennikeni sp. nov.; Neotrichia honda sp. nov.; Neotrichia landisae sp. nov.; Neotrichia lenati sp. nov.; Neotrichia mindyae sp. nov.; Neotrichia panamensis sp. nov.; Neotrichia parajarochita sp. nov.; Neotrichia paraxicana sp. nov.; Neotrichia snixae sp. nov.; Neotrichia spangleri sp. nov.; Neotrichia veraguasensis sp. nov. In addition, two new country records are presented: Neotrichia minutisimella (Chambers, 1873) and Neotrichia vibrans 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.
Aquatic insects, biodiversity, freshwater, Neotropics, protected areas
A concentrated effort during the last eight years (2015–2023) has almost doubled the known caddisfly fauna of Panama from 257 to 508 species distributed among 15 families and 56 genera (e.g.,
Before 2015, only three species of Neotrichia were known from Panama. Since then, we have added 25 new species and new country records (Table
Progression of Neotrichia species added to Panama’s caddisfly fauna beginning in 2015*. The left column indicates the sequence numbers in The Trichoptera of Panama publication series (citations below).
Trichoptera of Panama | Year | Number of new species | Number of new country records | Species |
---|---|---|---|---|
I | 2015 | 1 | Neotrichia canixa (Mosely, 1937) | |
II | 2015 | 2 | Neotrichia pamelae Harris & Armitage, 2015; Neotrichia parabullata Harris & Armitage, 2015 | |
IV | 2016 | 5 |
Neotrichia esmalda (Mosely, 1937); Neotrichia hiaspa (Mosely, 1937); Neotrichia tuxtla Bueno-Soria, 1999; Neotrichia unamas Botosaneanu (in |
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V | 2018 | 3 | Neotrichia anzuelo Armitage & Harris, 2018; Neotrichia collierorum Armitage & Harris, 2018; Neotrichia tatianae Armitage & Harris, 2018 | |
VI | 2018 | 1 | Neotrichia atopa Thomson & Armitage, 2018 | |
X | 2019 | 4 | Neotrichia carlsoni Harris & Armitage, 2019; Neotrichia rambala Harris & Armitage, 2019; Neotrichia serrata Harris & Armitage, 2019; Neotrichia starki Harris & Armitage, 2019 | |
2020 | 4 | Neotrichia espinosa Armitage & Harris, 2020; Neotrichia michaeli Armitage & Harris, 2020; Neotrichia pierpointorum Armitage & Harris, 2020; Neotrichia yayas Armitage & Harris, 2020 | ||
XVII | 2021 | 3 |
Neotrichia amplector Keth, 2004; Neotrichia armata Botosaneanu, 1993 (in |
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XX | 2023 | 2 | Neotrichia majagua Harris, Ríos & Aguirre, 2023; Neotrichia solapa Harris, Ríos & Aguirre, 2023 | |
XXIV | 2023 | 15 | 2 | New species or new country records in this publication: Neotrichia abrebotella sp. nov.; Neotrichia candela sp. nov.; Neotrichia codaza sp. nov.; Neotrichia embera sp. nov.; Neotrichia flennikeni sp. nov.; Neotrichia honda sp. nov.; Neotrichia landisae sp. nov.; Neotrichia lenati sp. nov.; Neotrichia mindyae sp. nov.; Neotrichia minutisimella (Chambers, 1873); Neotrichia panamensis sp. nov.; Neotrichia parajarochita sp. nov.; Neotrichia paraxicana sp. nov.; Neotrichia snixae sp. nov.; Neotrichia spangleri sp. nov.; Neotrichia veraguasensis sp. nov.; Neotrichia vibrans Ross, 1944 |
Totals: | 31 | 11 |
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.
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.
(Landis Reserve or Landis Reserva; Fig.
Maps A Panama, overlain by outlines of all 52 cuencas and showing all collection locations B collection locations (I–M) in the Pixvae area C collection locations (Q–T) in the Pajonal area. Key: A–Río Candela; B–Quebrada sin nombre; C–Quebrada sin nombre, locations 1 and 2; D–Quebrada sin nombre and Quebrada la Vuelta; E–Quebrada Grande; F–Quebrada Honda; G–Río Platanal; H–Quebrada San Cristobal; I–Quebrada Monita; J–Río Pixvae; K–Quebrada del Rosario; L–Quebrada La Mina; M–Quebrada El Rosario; N–Río Piedra de Moler; O–Quebrada Mulabá; P–Río Betegui; Q–Río Marica; R–Río Membrillo; S–Río Seren; T–Río Salado; U–Río Sajalices; V–Panama Canal; W–Quebrada sin nombre; X–Río Pirre. Note: Map colors are aesthetic, and do not impart any significance. Maps were generated using QGIS, v. 3.28.5-Firenze.
Landis Reserve is a private landholding in Chiriquí Province located north of Paso Canoas, which produces plants that are used in soil stabilization efforts. The stream sampled in this study is unnamed and of first order, located near the Costa Rican border. Ultimately, it flows into the Río Chiriquí Viejo and the Pacific Ocean. The riparian corridor is primarily forested with some open areas.
(Fig.
This is a large, private landholding in Chiriquí Province near San Andres bordered by the Río Cueta and the Río Gariché, both Pacific Ocean drainages. The immediate riparian corridor is forested, although this is surrounded by agricultural lands supporting a number of permanent crops such as cacao, plantains, and native tree species.
(PILA; Fig.
The Amistad International Park is a Latin American protected area shared by Panama and Costa Rica. Including a large portion of the Cordillera de Talamanca range, it encompasses approximately 401,000 ha of upland or montane tropical forest, and is an area of high diversity and endemicity.
(Fortuna Forest Reserve; Fig.
Part of Panama’s National System of Protected Areas, the Reserva Forestal Fortuna comprises 19,000 ha of primarily cloud forests, and an additional 500 ha of buffer zones. The 1,000 ha Fortuna Reservoir is a principal component of this protected area and is part of the Pacific drainage. The Reserva is also part of the Mesoamerican Biological Corridor, which includes the adjacent, and much larger, Bosque Protector Palo Seco (Caribbean Drainage) and Parque International La Amistad (Pacific and Caribbean drainage).
(Santa Fe NP or PNSF; Fig.
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 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.
(Altos de Campana NP or PNAC; Fig.
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.
(Darién NP or Darién National Park; Fig.
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.
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 1st to 3rd 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
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 (
Morphological terminology used for male genitalia generally follows that of
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 (
The genus Neotrichia (Hydroptilidae, Neotrichiinae) has a New World distribution with 211 species known from North, Central, and South America and the West Indies (
Panama: Chiriquí Province: Cuenca 102, Renacimiento District, Reserva Privada Landis, Location 1, Quebrada sin nombre; 8.643769°N, 82.829479°W; 755 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca 102, Renacimiento District, Reserva Privada Landis, Location 1, Quebrada sin nombre; 8.643769°N, 82.829479°W; 755 m a.s.l.; 15–31.iii.2020; M. Landis leg.; Malaise trap; MUPADI-003-T-2023 (in alcohol). Paratypes: Panama • 2 ♂♂; same as holotype; except, Location 2; 8.645005°N, 82.822037°W; 575 m a.s.l.; 27.ii–6.iii.2020; M. Landis leg.; Malaise trap; MUPADI-004-T-2023 (in alcohol).
Neotrichia abrebotella sp. nov. appears to be a member of the N. caxima group of
Male. Total length 1.7–1.9 mm (n = 3), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (Reserva Privada Landis).
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.
Panama: Chiriquí Province: Cuenca 102, Renacimiento District, Río Candela, Finca Félix, PILA; PSPSCB-PILA-C102-2017-021; 8.90614°N, 82.72882°W; 1799 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca 102, Renacimiento District, Río Candela, Finca Félix, PILA; PSPSCB-PILA-C102-2017-021; 8.90614°N, 82.72882°W; 1799 m a.s.l.; 1–5.ix.2017; E. Álvarez, T. Ríos, E. Pérez leg.; Malaise trap; MIUP-017-T-2023 (in alcohol).
Neotrichia candela sp. nov. is a member of the N. canixa group of
Male. Total length 1.7 mm (n = 1), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (Parque International La Amistad).
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.
Panama: Panama Oeste Province: Cuenca 115, Chame District, Altos de Campana NP, Río Sajalices; PSPSCB-PNAC-C115-2018-030; 8.67625°N, 79.89748°W; 194 m a.s.l.
Holotype : ♂, Panama: Panama Oeste Province: Cuenca 115, Chame District, Altos de Campana NP, Río Sajalices; PSPSCB-PNAC-C115-2018-030; 8.67625°N, 79.89748°W; 194 m a.s.l.; 29.v.2018; E. Pérez, C. Nieto, M. Molinar, T. Ríos leg.; UV light trap; MIUP-018-T-2023 (in alcohol).
Panama • ♂; Coclé Province: Cuenca 134, Penonomé District, Río Salado, Pajonal Geosite; 8.59580°N, 80.21512°W; 323 m a.s.l.; 4.iv.2022; C. Nieto leg.; UV light trap (in alcohol) • ♂; ibid., except Río Seren; 8.58983N, 80.21476W; 332 m a.s.l. • 2 ♂♂; ibid.., except Río Membrillo; 8.58450°N, 80.22074°W; 334 m a.s.l. • 17 ♂♂; ibid., except, Río Marica, Pajonal Geosite, 8.55016°N, 80.23831°W; 316 m a.s.l. • 2 ♂♂; Veraguas Province: Cuenca 116, Quebrada del Rosario; 7.85826°N, 81.55764°W; 26 m a.s.l.; 20.i.2023; V. Rodriguez leg.; UV light trap • 3 ♂♂; ibid., except Cuenca 132; Río Betegui;8.36047°N, 80.99481°W; 144 m a.s.l.; 28.i.2023; V. Rodriguez leg.; UV light trap.
Neotrichia codaza sp. nov. is a member of the N. collata group of
Male. Total length 1.3–1.5 mm (n = 12), 17 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Panama Oeste Province (Altos de Campana National Park).
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.
Panama: Darién Province: Cuenca 156, Pinogana District, Darién NP, Río Pirre, Estacion de MiAmbiente en Rancho Frio; 8.09081°N, 77.74043°W; 73 m a.s.l.
Holotype : ♂, Panama: Darién Province: Cuenca 156, Pinogana District, Darién NP, Río Pirre, Estacion de MiAmbiente en Rancho Frio; 8.09081°N, 77.74043°W; 73 m a.s.l.; 9–12.ii.2018; Malaise trap; A. Thurman leg.; MUPADI-005-T-2023 (in alcohol).
This new species, with a pair of apical phallic spines, would appear to be another member of the N. collata group of
Male. Total length 1.2 mm (n = 1), 17 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Darién Province (Darién National Park).
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.
Panama: Chiriquí Province: Cuenca 108, David District, San Pablo Viejo, puente vía Interamericana antes de llegar a la entrada de Bagala, Río Platanal; 8.46416°N, 82.52030°W; 825 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca 108, David District, San Pablo Viejo, puente vía Interamericana antes de llegar a la entrada de Bagala, Río Platanal; 8.46416°N, 82.52030°W; 825 m a.s.l.; 12.iv.2021; T. Ríos, Y. Aguirre leg.; UV light trap; MUPADI-006-T-2023 (in alcohol).
Neotrichia flennikeni sp. nov. appears to be a member of the N. vibrans group of
Male. Total length 1.8 mm (n = 1), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (David District).
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.
Panama: Chiriquí Province: Cuenca 108, Boquete District, Quebrada Honda, N Fortuna Dam, Fortuna Forest Reserve; 8.74985°N, 82.23885°W; 1132 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca 108, Boquete District, Quebrada Honda, N Fortuna Dam, Fortuna Forest Reserve; 8.74985°N, 82.23885°W; 1132 m a.s.l.; 18.ii.2018; B. Armitage, T. Arefina-Armitage leg.; UV light trap; MUPADI-007-T-2023 (in alcohol).
Neotrichia honda sp. nov. does not fit well in any of the species groupings established by
Male. Total length 1.5 mm (n = 1), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (Fortuna Forest Reserve).
This species is named for Quebrada Honda, a tributary of Río Chiriquí and Fortuna Reservoir in northeast Chiriquí Province, where the species was collected. The name is a noun in the genitive case.
Panama: Chiriquí Province: Cuenca 102, Renacimiento District, Reserva Privada Landis, Quebrada sin nombre, Location 2; 8.645005°N, 82.822037°W; 575 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca 102, Renacimiento District, Reserva Privada Landis, Quebrada sin nombre, Location 2; 8.645005°N, 82.822037°W; 575 m a.s.l.; 30.iv.2020; M. Landis leg.; Malaise trap; MUPADI-008-T-2023 (in alcohol). Paratypes: Panama • 9 ♂♂; ibid., except 30.v.2020; MUPADI-009-T-2023 (in alcohol).
Panama • 2 ♂♂; ibid., except 27.ii–15.iii.2020 • 1 ♂; ibid., except 15–31.iii.2020 • 2 ♂♂; ibid., except Location 1, 8.643769°N, 82.829479°W; 755 m a.s.l.; 27.ii–10.iii.2020; M. Landis leg.; Malaise trap • 1 ♂; Chiriquí Province: Bugaba District, Cuenca 102, nr San Andres, Finca La Esperanza, Quebrada La Vuelta; 8.61710°N, 82.70415°W; 492 m a.s.l.; 3–20.i.2022; T. Ríos, Y. Aguirre leg.; Malaise trap • 2 ♂♂; ibid., except 21.i–3.ii.2022 • 3 ♂♂; ibid., except 6–20.iii.2022 • 2 ♂♂; ibid., except 8–22.iv.2022 • 2 ♂♂; ibid., except 8–21.v.2022 • 2 ♂♂; ibid., except Quebrada sin nombre; 8.61765°N, 82.71330°W; 540 m a.s.l.; 21.i–3.ii.2022.
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 and N. 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.
Male. Total length 1.5–1.7 mm (n = 15), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (Reserva Privada Landis, Finca La Esperanza).
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.
Panama: Chiriquí Province: Cuenca102, Renacimiento District, Reserva Privada Landis, Location 1, Quebrada sin nombre; 8.643769°N, 82.82979°W; 755 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca102, Renacimiento District, Reserva Privada Landis, Location 1, Quebrada sin nombre; 8.643769°N, 82.82979°W; 755 m a.s.l.; 5–30.xii.2022; M. Landis leg.; Malaise trap; MUPADI-010-T-2023 (in alcohol).
Neotrichia lenati sp. nov. appears to be a member of the N. okopa group of
Male. Total length 1.4 mm (n = 1), 18 antennal segment, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (Reserva Privada Landis).
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.
Panama: Darién Province: Cuenca 156, Pinogana District, Darién NP, Río Pirre, Estacion de MiAmbiente en Rancho Frio; 8.09081°N, 77.74043°W; 73 m a.s.l.
Holotype : ♂, Panama: Darién Province: Cuenca 156, Pinogana District, Darién NP, Río Pirre, Estacion de MiAmbiente en Rancho Frio; 8.09081°N, 77.74043°W; 73 m a.s.l.; 9–12.ii.2018; A. Thurman leg.; Malaise trap; MUPADI-011-T-2023 (in alcohol).
Neotrichia mindyae sp. nov. is another member of the N. canixa group of
Male. Total length 1.5 mm (n = 1), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Darién Province (Darién National Park).
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.
Panama: Colon Province: Cuenca 117, Portobelo District, Quebrada sin nombre, nr Jose Pobre property–Tesoro Verde; 9.60069°N, 79.61658°W; 55 m a.s.l.
Holotype : ♂, Panama: Colon Province: Cuenca 117, Portobelo District, Quebrada sin nombre, nr Jose Pobre property–Tesoro Verde; 9.60069°N, 79.61658°W; 55 m a.s.l.; 19.xii.2018; D. Garrido leg.; UV light trap; MUPADI-012-T-2023 (in alcohol). Paratypes: Panama • 2 ♂♂; same as holotype; MUPADI-013-T-2023 (in alcohol).
Panama • 16 ♂♂; Veraguas Province: Cuenca 116, Las Palmas District, Quebrada La Mina; 7.87443°N, 81.51004°W; 63 m a.s.l.; 3.ii.2023; V. Rodríguez leg.; UV light trap • 7 ♂♂; ibid., except Río Indio; 7.87372°N, 81.49994°W; 57 m a.s.l.; 3.ii.2023 • 1 ♂; ibid., except Río Pixvae; 7.84287°N, 81.56329°W; 17 m a.s.l.; 23.i.2023 • 4 ♂♂; ibid., except Soná District, Quebrada Monita; 7.81480°N, 81.55724°W; 26 m a.s.l.; 21.i.2023.
Neotrichia panamensis sp. nov. is another member of the N. canixa group of
Male. Total length 1.5–1.7 mm (n = 14), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Colon Province (Portobelo District); Veraguas Province (Las Palmas District).
This species is named for the Republic of Panama where the species was collected. The name is a noun in the genitive case.
Panama: Chiriquí Province: Cuenca 102, Bugaba District, afluente Río Chiriquí Viejo, PILA; PSPSCB-PILA-C102-2017-022; 8.90124°N, 82.61817°W; 2354 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca 102, Bugaba District, afluente Río Chiriquí Viejo, PILA; PSPSCB-PILA-C102-2017-022; 8.90124°N, 82.61817°W; 2354 m a.s.l.; 17–21.vi.2017; E. Álvarez, E. Pérez, T. Ríos leg.; Malaise trap; MIUP-019-T-2023 (in alcohol).
Neotrichia parajarochita sp. nov. is another member of the N. canixa group of
Male. Total length 1.4 mm (n = 1), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (Parque International La Amistad).
This species is named for its resemblance to Neotrichia jarochita. The name is an adjective used as a substantive in the genitive case.
Panama: Veraguas Province: Cuenca 132, Santa Fe District, Santa Fe NP, Quebrada Mulabá, Santa Fe NP, PSPSCB-PNSF-C132-2017-009; 8.52560°N, 81.12956°W; 623 m a.s.l.
Holotype : ♂, Panama: Veraguas Province: Cuenca 132, Santa Fe District, Santa Fe NP, Quebrada Mulabá, Santa Fe NP, PSPSCB-PNSF-C132-2017-009; 8.52560°N, 81.12956°W; 623 m a.s.l.; 20.iv.2017; A. Cornejo, T. Ríos, C. Nieto leg.; UV light trap; MIUP-020-T-2023 (in alcohol).
Panama • ♂; Veraguas Province: Cuenca 097, Santa Fe District, Santa Fe NP, Río Piedra de Moler; PSPSCB-PNSF-C097-2017-012; 8.56553°N, 81.18817°W; 340 m a.s.l.; 20.iv.2017; A. Cornejo, T. Ríos, E. Álvarez, C. Nieto leg.; UV light trap • ♂; Panama Province: Panama Canal, date and locality of collection illegible on label; D. Denning leg.
Neotrichia paraxicana sp. nov. is another member of the N. canixa group of
Male. Total length 1.6–1.8 mm (n = 3), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Veraguas Province (Santa Fe National Park).
This species is named for its overall resemblance to Neotrichia xicana. The name is an adjective used as a substantive in the genitive case.
Panama: Chiriquí Province: Cuenca 102, Renacimiento District, Reserva Privada Landis, Quebrada sin nombre, Location 1; 8.64379°N, 82.82949°W; 755 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca 102, Renacimiento District, Reserva Privada Landis, Quebrada sin nombre, Location 1; 8.64379°N, 82.82949°W; 755 m a.s.l.; 15–31.iii.2020; M. Landis leg.; Malaise trap; MUPADI-014-T-2023 (in alcohol).
Panama • ♂; Chiriquí Province: Cuenca 108, Boquete District, Quebrada Jaramillo Abajo; 8.745827°N, 82.418083°W; 1054 m a.s.l.; 6.ii.2019; K. Castillo leg.; UV light trap.
Neotrichia snixae sp. nov. is a member of the N. canixa group of
Male. Total length 1.9 mm (n = 2), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (Reserva Privada Landis; Boquete District).
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.
Panama: Chiriquí Province: Cuenca 108, Boquete District, Bajo Boquete, Quebrada Cheche, Hotel Fundadores; 8.77195°N, 82.43308°W; 1200 m a.s.l.
Holotype : ♂, Panama: Chiriquí Province: Cuenca 108, Boquete District, Bajo Boquete, Quebrada Cheche, Hotel Fundadores; 8.77195°N, 82.43308°W; 1200 m a.s.l.; 29.v.1983; P. Spangler, R. Faitoule, W. Steiner leg.; MUPADI-015-T-2023 (in alcohol). Paratype. Panama • ♂; same as holotype; MUPADI-016-T-2023 (in alcohol).
Panama • 2 ♂♂; Chiriquí Province: Cuenca 104, Bugaba District, La Concepción, Río Guigala, Puente antiqua vias del Ferrocarril; 8.51845°N, 82.64280°W; 209 m a.s.l.; 12.iii.2021; T. Ríos, Y. Aguirre leg.; UV light trap • 2 ♂♂; ibid., except Cuenca 108, Boquerón District, Río Chirigagua, Puente antes de llegar al Hotel Los Delfines; 8.48139°N, 82.54788°W; 128 m a.s.l.; 12.iv.2021; T. Ríos, Y. Aguirre leg.; UV light trap • 4 ♂♂; ibid., except David District, San Pablo Viejo, puente vía Interamericana antes de llegar a la entrada de Bagala, Río Platanal; 8.46416°N, 82.52030°W; 84 m a.s.l.; 12.ii.2021; T. Ríos, Y. Aguirre leg.; UV light trap • ♂; ibid., except 12.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).
The pair of spines at the phallic apex and the posterolateral process from segment IX places this species in the N. collata group of
Male. Total length 1.3–1.5 mm (n = 10), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Chiriquí Province (Boquerón, Boquete, and Bugaba districts); Darién Province (Chepigana District).
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.
Panama: Veraguas Province: Cuenca 132, Santa Fe District, Santa Fe NP, Quebrada Mulabá; PSPSCB-PNSF-C132-2017-009; 8.52560°N, 81.12956°W; 623 m a.s.l.
Holotype : ♂, Panama: Veraguas Province: Cuenca 132, Santa Fe District, Santa Fe NP, Quebrada Mulabá; PSPSCB-PNSF-C132-2017-009; 8.52560°N, 81.12956°W; 623 m a.s.l.; 20.iv.2017; A. Cornejo, T. Ríos, E. Álvarez, C. Nieto leg.; UV light trap; MIUP-021-T-2023 (in alcohol).
Panama • ♂; ibid., except Río Piedra de Moler; PSPSCB-PNSF-C097-2017-012; 8.56553°N, 81.18817°W; 340 m a.s.l.; 20.iv.2017; MIUP (in alcohol).
This new species, which lacks spines at the phallic apex, is placed in the N. okopa group of
Male. Total length 1.6 mm (n = 2), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Panama: Veraguas Province (Santa Fe National Park).
This species is named for Veraguas Province where the male holotype was collected. The name is a noun in the genitive case.
Panama • ♂; Veraguas Province: Cuenca 097, PNSF, afluente sin nombre de Río Calovebora; PSPSCD-PNSF-CO97-2017-011; 8.55343°N, 81.17675°W; 395 m a.s.l.; 20.iv.2017; A. Cornejo, T. Ríos, E. Álvarez, C. Nieto leg.; UV light trap.
This is a new country record for Panama and a significant extension of its southern range (formerly Texas, U.S.A.).
Canada: Manitoba; Panama: Veraguas Province (Santa Fe National Park); U.S.A.: Alabama, Arkansas, Florida, Georgia, Illinois, Indiana, Kansas, Kentucky, Louisiana, Minnesota, Mississippi, Missouri, North Carolina, Oklahoma, South Carolina, Texas.
Panama • 27 ♂♂, 25 ♀♀; Chiriquí Province: Cuenca 108, David, UNACHI–Jardin Botanico, El Cabrero, nr Quebrada San Cristobal; 8.434060°N, 82.451930°W; 45 m a.s.l.; 19.iv–3.v.2021; Y. Aguirre, T. Ríos leg.; Malaise trap • 22 ♂♂, 23 ♀♀; ibid., except 5–19.iv.2021 • ♂; Panama Oeste Province: Cuenca 115, Chame District, Altos de Campana NP, Río Sajalices; PSPSCB-PNAC-C115-2018-030; 8.67625°N, 79.89748°W; 194 m a.s.l.; 27–31.v.2018; E. Pérez, C. Nieto, M. Molinar, T. Ríos leg.; Malaise trap.
Female (Fig.
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.
Females of other species from the N. vibrans group of
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
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.
Mexico: Chihuahua; Panama: Chiriquí Province (David District); U.S.A.: Alabama, Arkansas, Florida, Georgia, Illinois, Kansas, Kentucky, Maine, Minnesota, Mississippi, Missouri, New Hampshire, New York, Ohio, Oklahoma, South Carolina, Tennessee, Texas, Virginia, West Virginia, Wisconsin.
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.
1 | Inferior appendages fused mesally into a plate, which is evident in ventral view (Fig. |
2 |
– | Inferior appendages separate, not fused mesally as seen in ventral view (Figs |
3 |
2 | Inferior appendages fused into an elongate rectangular plate ( |
N. minutisimella (Chambers, 1873) |
– | Inferior appendages fused into an ovate plate (Fig. |
N. vibrans Ross, 1944 |
3 | Horns from posterior tergum X, which vary in shape and size (Figs |
4 |
– | Horns absent from posterior tergum X (Figs |
20 |
4 | Horns from posterior tergum X asymmetrical (Figs |
5 |
– | Horns from posterior tergum X symmetrical (Figs |
7 |
5 | Left-side horn of posterior tergum X longer than right-side horn (Fig. |
N. parajarochita sp. nov. |
– | Right-side horn of posterior tergum X longer than left-side horn (Fig. |
6 |
6 | Posterolateral process from segment IX present; ventral branch of bracteole longer than dorsal branch (Fig. |
N. snixae sp. nov. |
– | Posterolateral process from segment IX absent ( |
N. malickyi Harris, 1993 (in |
7 | Linear posterior process from segment IX present (Fig. |
8 |
– | Linear posterior process from segment IX absent ( |
12 |
8 | Linear posterior process from segment IX short, not extending beyond horns in lateral view (Fig. |
N. paraxicana sp. nov. |
– | Linear posterior process from segment IX elongate, extending beyond horns in lateral view ( |
9 |
9 | Linear posterior process from segment IX serrate ( |
10 |
– | Linear posterior process from segment IX not serrate, or lobate or cleft distally (Fig. |
11 |
10 | Linear posterior process from segment IX in lateral view serrate on dorsal margin ( |
N. serrata Harris & Armitage, 2019 |
– | Linear posterior process from segment IX not serrate on distal margin, cleft or lobate distally ( |
N. xicana (Mosely, 1937) |
11 | Linear posterior process from segment IX widening distally and setose on margin ( |
N. canixa (Mosely, 1937) |
– | Linear posterior process from segment IX saber-like, narrowing distally and without setose margin ( |
N. unamas Botosaneanu, 1993 (in |
12 | Dorsal branch of bracteole shorter than ventral branch ( |
N. tauricornis Malicky, 1980 |
– | Dorsal branch of bracteole longer than ventral branch, which may be vestigial (Figs |
13 |
13 | Bracteole with ventral branch ≥ ½ as long as dorsal branch (Fig. |
14 |
– | Bracteole with ventral branch < ½ as long as dorsal branch and in some cases vestigial (Figs |
17 |
14 | Inferior appendage in lateral view with dorsal hump at midlength ( |
N. collierorum Armitage & Harris, 2018 |
– | Inferior appendage in lateral view parallel-sided ( |
15 |
15 | Phallus apex undivided, narrowing to elongate slender process ( |
N. anzuelo Armitage & Harris, 2018 |
– | Phallus apex divided ( |
16 |
16 | Inferior appendage in lateral view saber-shaped ( |
N. majagua Harris, Ríos & Aguirre, 2023 |
– | Inferior appendage in lateral view cigar-shaped (Fig. |
N. candela sp. nov. |
17 | Phallus apex with single elongate process with small notch near base (Fig. |
18 |
– | Phallus apex clearly divided into two processes (Fig. |
19 |
18 | Posterior horns from tergum X strongly curving inward (Fig. |
N. mindyae sp. nov. |
– | Posterior horns from tergum X not strongly curving inward ( |
N. michaeli Armitage & Harris, 2020 |
19 | Dorsal branch of inferior appendage in lateral view curving and widening apically ( |
N. pamelae Harris & Armitage, 2015 |
– | Dorsal branch of inferior appendage in lateral view not curving and widening sub-basally (Fig. |
N. panamensis sp. nov. |
20 | Posterolateral process from segment IX, which takes different forms, from elongate (Fig. |
21 |
– | Posterolateral process from segment IX absent (Fig. |
36 |
21 | Posterolateral process from segment IX elongate, exceeding length of inferior appendage and bracteole, heavily sclerotized and narrowing distally to an acute point (Fig. |
22 |
– | Posterolateral process from segment IX shorter, typically not exceeding length of inferior appendage and bracteole (Fig. |
26 |
22 | Inferior appendage in lateral view wide basally, greatly tapering distally to uniform width (fig. 13A in |
N. hiaspa (Mosely, 1937) |
– | Inferior appendage various but not wide basally and tapering distally (Fig. |
23 |
23 | Elongate, thin ventroposterior process from segment XI; subgenital plate in lateral view with elongate ventral process ( |
N. yayas Armitage & Harris, 2020 |
– | Elongate, thin, ventroposterior process from segment IX absent (Fig. |
24 |
24 | Posterolateral processes from segment IX nearly symmetrical in dorsal view (Fig. |
25 |
– | Posterolateral processes from segment IX asymmetrical in dorsal view ( |
N. tatianae Armitage & Harris, 2018 |
25 | Posterolateral process from segment IX sinuate in lateral and dorsal view ( |
N. starki Harris & Armitage, 2019 |
– | Posterolateral process from segment IX linear in lateral and dorsal views (Fig. |
N. landisae sp. nov. |
26 | Posterolateral process from segment IX spinose, heavily sclerotized and tapering distally (Fig. |
27 |
– | Posterolateral process from segment IX various, but not spinose of heavily sclerotized (Figs |
32 |
27 | Phallus apex divided into pair of elongate spines (Fig. |
28 |
– | Phallus apex not divided into pair of elongate spines ( |
29 |
28 | Segment IX in lateral view with elongate mesal process ( |
N. pierpointorum Armitage & Harris, 2020 |
– | Segment IX in lateral view without elongate mesal process ( |
N. esmalda (Mosely, 1937) |
29 | Inferior appendage in lateral and ventral views elongate and thin, not heavily sclerotized ( |
30 |
– | Inferior appendage in lateral and ventral views short and triangular, heavily sclerotized Fig. |
N. honda sp. nov. |
30 | Two posterolateral sclerotized processes from segment IX ( |
N. espinosa Armitage & Harris, 2020 |
– | Single posterolateral process from segment IX ( |
N. carlsoni Harris & Armitage, 2019 |
31 | Phallus with pair of sclerotized spines (Figs |
32 |
– | Phallus with single sclerotized spine (Fig. |
34 |
32 | Phallic apical spines terminal (Figs |
33 |
– | Phallic apical spines subterminal (Fig. |
N. codaza sp. nov. |
33 | Phallic apical spines short and nearly equal in length (Fig. |
N. embera sp. nov. |
– | Phallic apical spines elongate and subequal in length (Fig. |
N. spangleri sp. nov. |
34 | Phallus with sclerotized spine (Fig. |
35 |
– | Phallus without sclerotized spine (Fig. |
N. lenati sp. nov. |
35 | Posterolateral process from segment IX widening distally and serrate (fig. 5A, B in |
N. solapa Harris, Ríos & Aguirre, 2023 |
– | Posterolateral process from segment IX not widening distally or serrate (Fig. |
N. flennikeni sp. nov. |
36 | Inferior appendages in lateral view with serrate dorsal process just past midlength ( |
N. parabullata Harris & Armitage, 2015 |
– | Inferior appendages various but never with a serrate dorsal process just past midlength (Figs |
37 |
37 | Inferior appendages short in lateral view, not extending much beyond subgenital plate or bracteoles (Fig. |
38 |
– | Inferior appendages elongate in lateral view, typically extending well beyond the subgenital plate and bracteoles (Fig. |
41 |
38 | Bracteole in lateral view large and nearly circular in shape ( |
N. atopa Thomson & Armitage, 2018 |
– | Bracteole in lateral view thin over length, not nearly circular (Fig. |
39 |
39 | Inferior appendage in ventral view terminating in elongate spines ( |
40 |
– | Inferior appendage in ventral view not terminating in elongate or short spines (Fig. |
N. abrebotella sp. nov. |
40 | Inferior appendage in ventral view terminating in elongate spines (fig. 76C in |
N. armata Botosaneanu, 1993 (in |
– | Inferior appendages in ventral view terminating in short spines ( |
N. amplector Keth, 2004 |
41 | Phallus terminating in a spine ( |
42 |
– | Phallus not terminating in a spine or spines (Fig. |
44 |
42 | Phallus terminating in a single spine ( |
N. rambala Harris & Armitage, 2019 |
– | Phallus terminating in a pair of rods which can be fine ( |
43 |
43 | Rods at apex of phallus stout and elongate ( |
N. tuxtla Bueno-Soria, 1999 |
– | Rods at apex of phallus thin and short ( |
N. flowersi Harris, 1990 |
44 | Inferior appendages short and triangular in lateral view (Fig. |
N. veraguasensis sp. nov. |
– | Inferior appendages elongate and rectangular in lateral and ventral views ( |
N. kampa Oláh & Johanson, 2011 |
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 (
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
Whereas the majority of the new species herein described derived from field activities of AIRG at MUPADI, six of the new species and specimen records included herein were acquired through the Sustainable Production System and Biodiversity Conservation Project (PSPSCB), conducted in a variety of Panamanian national parks and protected areas. We acknowledge the Panamanian Ministry of Environment, which managed this project with funding from the World Bank, and who provided collecting permits and other support. We thank Yusseff P. Aguirre, Eric Álvarez, Milexi Molinar, Carols Nieto, Edgar Pérez, and Aydeé Cornejo who collectively were involved in all or some of the national park collections. We also appreciate the organizational and logistical support by the Gorgas Institute and COZEM concerning this project. Collections from the Pajonal area of Panama (Fig.
The authors have declared that no competing interests exist.
No ethical statement was reported.
We acknowledge the Panamanian Ministry of Environment, which managed the Sustainable Production System and Biodiversity Conservation Project with funding from the World Bank.
Conceptualization: BJA. Investigation: TARG, SCH. Data curation: BJA, TARG. Formal analysis: SCH. Writing – original draftt: BJA, SCH. Writing–Review and editing: BJA, SCH, TARG. Funding acquisition: BJA.
Steven C. Harris https://orcid.org/0000-0002-6432-7462
Brian J. Armitage https://orcid.org/0000-0003-3182-1533
Tomás A. Ríos González https://orcid.org/0000-0003-0590-6488
All of the data that support the findings of this study are available in the main text.