Research Article |
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Corresponding author: Brian J. Armitage ( tobikera89@gmail.com ) Academic editor: Gleison Desiderio
© 2025 Brian J. Armitage, Steven C. Harris, Viterbo Rodriguez.
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:
Armitage BJ, Harris SC, Rodriguez V (2025) The Trichoptera of Panama XXVIII. Nine new species of microcaddisflies (Trichoptera, Hydroptilidae). Neotropical Biology and Conservation 20(2): 125-147. https://doi.org/10.3897/neotropical.20.e148738
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In this study, we describe and illustrate nine new species of microcaddisflies: Oxyethira jaramillo sp. nov., Neotrichia afuera sp. nov., Neotrichia aguirrei sp. nov., Neotrichia betegui sp. nov., Neotrichia rancheria sp. nov., Neotrichia sona sp. nov., Metrichia riosi sp. nov., Ochrotrichia mariettae sp. nov. and Ochrotrichia martinez sp. nov. The majority of these new species were collected on the Soná Peninsula or offshore on the Coiba National Park islands. With these additions, there are now 544 species of caddisflies known from Panama, distributed amongst 15 families and 56 genera.
Neotropics, Isla Canales de Afuera, Isla Rancheria, Coiba National Park
During the last nine years, a concentrated investigative effort has more than doubled the number of known caddisfly species for Panama from 257 in 2015 to 535 in 2024, distributed amongst 15 families and 56 genera (
The Aquatic Invertebrate Research Group (AIRG) at the Universidad Autónoma de Chiriquí (
A comprehensive summary of the results of AIRG research (2015 through 2024) into the Trichoptera of Panama can be found in
The flora and fauna of Panama is species-rich due to its tropical location and topography and the resulting diversity of microclimates and habitats. In terms of aquatic habitats, Panama contains over 500 rivers, countless creeks and other types of lotic and lentic waterbodies. A more thorough description of its geology, topography, climate and habitats can be found in
The Jaramillo Corregimiento is a political subdivision (77.5 km2) of Boquete District in Chiriqui Province, Panama. Located west of Boquete, it is the first range east of Volcan Baru which anchors the western end of the Central Cordillera of Panama. Primarily forested and mountainous, the hydrography on its south-western face includes Quebrada Jaramillo and its tributaries.
Maps of collection sites A Panama map with administrative unit (provinces and indigenous areas) overlay showing the relative positions of all sampling locations pertinent to this publication B expanded view of the western Soná Peninsula and northern portion of Coiba National Park C Panama map with a major cuenca (watershed) overlay showing the location of collection sites for Ochrotrichia martinez sp.nov., the most widely distributed species within Panama of the nine species described herein [A–Landis Reserve; B–Quebrada Jaramillo Abajo; C–Quebrada Martínez; D–Río Calovebora; E– Quebrada Primo Brazo Mulabá; F–Quebrada Corazones; G–Río Beteguí; H–Isla Rancheria; I–Isla Canales de Afuera; J–Quebrada Monita; K–Quebrada del Rosario; L–Quebrada la Mina; M–Río Indio; N–Río Limón].
Landis Reserve is a private landholding in Chiriquí Province located north of Paso Canoas, which produces plants that are used in soil stabilisation 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 from there into the Pacific Ocean. The riparian corridor is primarily forested with some open areas.
Coiba National Park is located in the Gulf of Chiriqui off Panama’s Pacific coast, south of the Soná Peninsula and west of the Azuero Peninsula, the latter defining the eastern limit of the Gulf. The Park is a marine reserve that incorporates or includes 38 islands and the waters surrounding them (in total, almost ~ 70,585 acres). In 2005, the Park was recognised as a World Heritage Site by UNESCO.
Soná Peninsula is an unofficial name for a portion of Panama’s southern coast that juts out into the Gulf of Chiriqui (Pacific Ocean). It is comprised of the Soná District and Soná Corregimiento of Veraguas Province. It is immediately west of the much larger Azuero Peninsula. These two peninsulas and the nearby Coiba Island include accreted terrains transported in the past by the Pacific plate as it underwent subduction under the Caribbean plate and incorporated into Panama’s landmass.
The Bosque Protector Palo Seco protected area in western Panama occupies a portion of the Talamanca Cordillera, Central Cordillera and Caribbean slope lowlands and is part of both the Ngäbe-Buglé Comarca and Bocas del Toro Province. It connects the eastern part of La Amistad International Park with the Fortuna Forest Reserve. The Río Guabo watershed is a dendritic, Caribbean watershed, with a first-order creek, Quebrada Martınéz. The Río Guabo empties into the Laguna de Chiriquí, part of the Caribbean Sea, immediately west of the port of Chiriquí Grande. The landscape through which it flows is primarily forested with some agriculture and with a forested riparian corridor for at least some of its length.
National parks and forest reserves indicated above are under the stewardship of the Ministerio de Ambiente (MiAmbiente) and are 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 forested riparian corridors.
The majority of the collections made in alcohol by AIRG employed light traps (UV fluorescent tubes or an UV LED strips). Whenever possible, Malaise traps were also set up in the same locations, because the combination of UV light traps and Malaise traps has consistently produced the best results in terms of species diversity. Sampling monthly from December to June has produced very good results, equivalent to whole-year sampling. Collecting just once at a given location produces insufficient information about the species assemblage present. Once again, optimum results are achieved with multiple-month collections at the same site and with the two collection methods stated above. Specimens were prepared and examined following standard methods outlined in
Morphological terminology used for male genitalia generally follows that of
Holotypes listed in this publication are deposited in Museo de Peces de Agua Dulce e Invertebrados (MUPADI) at the Universidad Autónoma de Chiriquí. Paratypes and other specimens are deposited in MUPADI, the University of Minnesota’s Neotropical Insect Collection (
Order Trichoptera Kirby, 1813
Suborder Integripalpia Martynov, 1924
Superfamily Hydroptiloidea Stephens, 1836
Family Hydroptilidae Stephens, 1836
The genus Oxyethira includes over 253 species of caddisflies (
Panama: Chiriqui Province, Cuenca 108, Boquete District, Quebrada Jaramillo Abajo; 8.745827°N, 82.418083°W, 1059 m a.s.l.
Holotype • ♂, in alcohol. Original label: “Panama: Chiriqui Province, Cuenca 108, Boquete District, Quebrada Jaramillo Abajo; 8.745827°N, 82.418083°W, 1059 m a.s.l.; 9 Mar 2019; leg K. Castillo; UV light trap"; MUPADI.
Oxyethira jaramillo sp. nov. is related to a number of species which have elongate dorsal processes from segment IX, such as O. bidentata (Mosely), O. culebra Holzenthal & Harris and other members of the O. aeola group (
Male. Total length 2.8 mm (n = 1), 29 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named after both the type waterbody, Quebrada Jaramillo, in the western Panamanian highlands and the reversal of the earth’s polarity ~ 900,000 years ago, referred to as the Jaramillo Event (
Subfamily Neotrichiinae Ross, 1956
The genus Neotrichia, found in North, Central and South America and the West Indies, contains over 228 species (
Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.
Holotype • ♂, in alcohol. Original label: “Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.; 6–22 Mar 2023; leg V. Rodriguez; Malaise trap”; MUPADI.
Neotrichia afuera sp. nov. does not fit well within the species groups established by
Male. Total length 1.4 mm (n = 1), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named for the isle in Panama where the species was collected.
Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.
Holotype • ♂, in alcohol. Original label: “Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.; 6–22 Mar 2023; leg V. Rodriguez; Malaise trap”; MUPADI. Paratype • ♂, in alcohol; same as holotype.
♂, in alcohol; Panama, Veraguas Province, Cuenca 116, Soná District, Río Limon; 7.87182°N, 81.49397°W; 67 m a.s.l.; 18 Apr 2023; leg V. Rodriguez; UV light trap; MUPADI.
Neotrichia aguirrei sp. nov. is a member of the N. canixa group of
Male. Total length 1.8 mm (n = 3), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named for Yusseff P. Aguirre Espinoza, Panamanian biologist at the Museo de Peces de Agua Dulce e Invertebrados of the Universidad Autónoma de Chiriquí in recognition of his intensive and extensive involvement with our studies of aquatic insects in Panama. The name is a noun in the genitive case.
Panama: Veraguas Province, Cuenca 132, San Francisco District, nr. La Perdiz, N of San Francisco, Río Beteguí; 8.36047°N, 80.99481°W, 144 m a.s.l.
Holotype • ♂, in alcohol. Original label: “Panama: Veraguas Province, Cuenca 132, San Francisco District, nr. La Perdiz, N of San Francisco, Río Beteguí; 8.36047°N, 80.99481°W, 144 m a.s.l.; 28 Jan 2023; leg V. Rodriguez; UV light trap”; MUPADI.
Neotrichia betegui sp. nov. appears to be a member of the Neotrichia caxima group of
Male. Total length 1.7 mm (n = 1), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named for the Río Beteguí from which the type specimen was collected. The derivation of "betegui" is unclear. The prefix "be" means exists and the Aztec word for lizard translates into Spanish as "tegui". However, we have no confirmation that this was the genesis of the specific epithet. The name is a noun in the genitive case.
Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.
Holotype
• ♂, in alcohol. Original label: “Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.; 6–22 Mar 2023; leg V. Rodriguez; Malaise trap”; MUPADI. Paratypes • 9 ♂♂, in alcohol; same as holotype; MUPADI • 1 ♂; ibid., Isla Rancheria; 7.6396°N, 81.70475°W; 24 m a.s.l.; 6–21 Mar 2023; MUPADI (7 ♂♂),
The Neotrichia canixa group consists of a cluster of Neotropical species with an elongate dorsal process from segment IX, including N. filifera Flint and N. napoensis Harris & Davenport, but the new species is closest to N. starki Harris & Armitage from Bocas del Toro Province in Panama. Neotrichia rancheria sp. nov. is separated from N. starki by the shorter rods from segment XI, the setose subgenital plate and the closely appressed inferior appendages.
Male. Total length 1.2–1.4 mm (n = 10), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named for the Pacific island, Isla Ranchera, off the southern coast of Panama where the type specimens were collected. This island is part of Coiba National Park, which is a UNESCO World Heritage Site and Marine Park. Isla Rancheria, also known as Isla Coibita, is located off the NE coast of Coiba Island. The name is a noun in the genitive case.
Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.
Holotype
• ♂, in alcohol. Original label: Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.; 20 Jan–5 Feb 2023; leg V. Rodriguez; Malaise trap”; MUPADI. Paratypes • 3 ♂♂, in alcohol; same as holotype, except:
4 ♂♂, in alcohol; ibid., Cuenca 116, Las Palmas District, Quebrada del Rosario; 7.85826°N, 81.55764°W; 26 m a.s.l.; 3 Feb 2023; leg V. Rodriguez; Malaise trap; MUPADI • 1 ♂; ibid., 20 Jan–21 Feb 2023; Malaise trap; MUPADI • 3 ♂♂; ibid., 20 Jan 2023; UV light trap; MUPADI • 1 ♂; ibid., Río Indio; 7.8710°N, 81.49459°W; 703 m a.s.l.; 3 Feb 2023; MUPADI • 4 ♂♂; ibid., Quebrada La Mina; 7.87443°N, 81.51004°W; 63 m a.s.l.; MUPADI • 3 ♂♂; ibid., Soná District, Quebrada Monita; 7.81480°N, 81.55724°W; 26 m a.s.l.; 20 Jan–4 Feb 2023; Malaise trap; MUPADI • 15 ♂♂; ibid., 21 Jan 2023, UV light trap; MUPADI • 17 ♂♂; ibid., 11 Apr 2023;
Neotrichia sona sp. nov. appears to be a member of the Neotrichia vibrans group of
Male. Total length 1.6–1.8 mm (n = 30), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named for the peninsula, district, corregimiento and town along the Pacific coast of Panama. One suggested derivation of the word "soná" is from a local indigenous language, in which sonáre means “murmur of the waters”. This refers to the numerous riverbeds that surround the region. The name is a noun in the genitive case.
Subfamily Ochrotrichiinae Marshall, 1979
The genus Metrichia includes over 141 species of caddisflies (
Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.
Holotype • ♂, in alcohol. Original label: Panama: Veraguas Province, Soná District, Canales de Afuera Island off the Soná Peninsula nr. Pixvae; 7.69494°N, 81.62649°W; 29 m a.s.l.; 6–22 Mar 2023; leg V. Rodriguez; Malaise trap”; MUPADI. Paratypes • 5 ♂♂, in alcohol; same as holotype; MUPADI.
Metrichia riosi sp. nov. is similar to M. trebeki Harris & Armitage, M. rawlinsi Flint & Sykora and M. triquetra Bueno-Soria & Holzenthal, based on the shape of inferior appendage in lateral view, but it differs from these species in the structure of the phallus, which has three subapical spines.
Male. Total length 2.1–2.3 mm (n = 6), 18 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named for Tomás A. Ríos González, Panamanian biologist at the Museo de Peces de Agua Dulce e Invertebrados of the Universidad Autónoma de Chiriquí in recognition of his intensive and extensive involvement with our studies of aquatic insects in Panama. The name is a noun in the genitive case.
The genus Ochrotrichia includes over 226 species of caddisflies (Thompson 2023) distributed primarily in North, Central and South America. Currently, there are 42 species known from Panama (
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 • ♂, in alcohol. Original label: 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.; 30 Dec 2022; leg M. Landis; Malaise trap”; MUPADI. Paratype • ♂, ibid., same as holotype, except Location 2; 8.64501°N, 82.82204°W; 575 m a.s.l.; 29 Feb–16 Mar 2020; MUPADI.
Based on the sigmoid shape of the inferior appendage which bears thick spines apically, this species is placed in the O. arranca group of
Male. Total length 2.0 mm (n = 2), 26 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named in honour of Señora Marietta Landis, who provided permission and access to her private reserve in western Panama near the Costa Rican border and who made all of the collections, including those of this and several other new species. The name is a noun in the genitive case.
Panama: Comarca Ngäbe-Buglé, Cuenca 093, Kankintú District, Bosque Protector Palo Seco, Quebrada Martínez, detrás de las caseta de MiAmbiente; 8.79484°N, 82.19047°W; 480 m a.s.l.
Holotype • ♂, in alcohol. Original label: “Panama: Comarca Ngäbe-Buglé, Cuenca 093, Kankintú District, Bosque Protector Palo Seco, Quebrada Martínez, detrás de las caseta de MiAmbiente; 8.79484°N, 82.19047°W; 480 m a.s.l.; 12–29 Nov 2019, leg Y. Aquirre and T. Ríos; Malaise trap”; MUPADI. Paratypes • 3 ♂♂, in alcohol, ibid., 13–27 Apr 2019; MUPADI • 4 ♂♂, in alcohol, ibid., 24 Apr–8 May 2019; MUPADI • 12 ♂♂, in alcohol, ibid., 24 May–9 Jun 2019; MUPADI • 4 ♂♂, in alcohol, ibid., 8–22.vi.2019; MUPADI • 2 ♂♂, in alcohol, ibid.,16–30 Aug 2019; MUPADI • 1 ♂, in alcohol, ibid.,13–27 Sep 2019;
21 ♂♂, in alcohol, Cocle Province, Cuenca 105, La Pintada District, Omar Torrijos National Park, Quebrada Corazones, PSPSCBPNGDOTH-C103-2017-001, 8.67760°N, 80.60007°W, 728 m a.s.l.; 24 Mar 2017; leg A. Cornejo, E. Pérez, T. Ríos, E. Alvarez, and C. Nieto; UV light trap • 1 ♂, in alcohol, ibid., afluente Quebrada Corazones, PSPSCBPNGDOTH-C103-2017-002; 8.67801°N, 80.60006°W, 792 m a.s.l.; 24 Mar 2017; leg A. Cornejo, T. Ríos, E. Álvarez, and E. Pérez; UV light trap • 4 ♂♂, in alcohol, Chiriqui Province, Cuenca 108, Boquete District, Quebrada Jaramillo Abajo; 8.74583°N, 82.41808°W, 1059 m a.s.l.; 6 Mar 2019; UV light trap; leg K. Castillo • 5 ♂♂, in alcohol, Comarca Ngäbe-Buglé, Cuenca 093, Kankintú District, Bosque Protector Palo Seco, Quebrada Martínez, Willie Mazu; 8.79361°N, 82.19391°W, 538 m a.s.l.; 30 Aug 2019; leg Y. Aquirre and T. Ríos; UV light trap; 6 ♂♂, in alcohol, ibid.,16–30 Aug 2019; Malaise trap • 2 ♂♂, in alcohol, ibid.,13–27 Sep 2019 • 5 ♂♂, in alcohol, ibid., 27 Sep–11 Oct 2019 • 4 ♂♂, in alcohol, ibid., 8–12 Oct 2019 • 6 ♂♂, in alcohol, ibid.,11–30 Oct 2019 • 1 ♂, in alcohol, ibid.,12–26 Nov 2019 • 3 ♂♂, in alcohol, ibid.,12–29 Dec 2019 • 21 ♂♂, in alcohol, Veraguas Province, Cuenca 097, Santa Fe District, Santa Fe National Park, Río Calovebora, PSPSCD-PNSF-CO97-2017-005; 8.54318°N, 81.16398°W; 536 m a.s.l., 19–23 Apr 2017, leg T. Ríos, E. Álvarez and C. Nieto; Malaise trap • 28 ♂♂, in alcohol, ibid., Cuenca 132, Quebrada Primo Brazo Mulabá, PSPSCD-PNSF-C132-2017-008; 8.51706°N, 81.12181°W; 770 m a.s.l.; 19 Apr 2017.
Based on the elongate inferior appendages, this species is placed in the Ochrotrichia tenango group of
Male. Total length 2.5–3.5 mm (n = 25), 38 antennal segments, wings and body brown in alcohol. Genitalia (Fig.
Unknown.
Named for the stream, Quebrada Martínez, in which the types were collected. Located in the Bosque Protector Palo Seco Protected Area, administered by the Ministerio de Ambiente, on the Panama’s Caribbean slope, this is a first order stream in the Río Guabo watershed of the Ngäbe Buglé Comarca (Kankintú District). The word Martínez is a masculine name peculiar to Hispanic-American communities and means “son of Martin”. “Martin” is the Spanish form of the name “Martinus”, associated with the name Mars, which from the Latin implies “male”. The name is a noun in the genitive case.
Initially, our collections on the Soná Peninsula were made to compare and contrast with caddisfly assemblages on the islands of Coiba National Park. So far, for all Trichoptera species (unpublished data), there appears to be some overlap between these two areas, but also more than a few unique species on the islands. All of the five new species above from these two areas (Neotrichia afuera sp. nov., N. aguirrei sp. nov., N. rancheria sp. nov., N. sona sp. nov. and Metrichia riosi sp. nov.) were described from Isla Canales de Afuera, an island ~ 6 km distant from the mainland. One of these five species, Neotrichia rancheria was also found on Isla Rancheria (~ 16 km distant from the mainland and very close to Coiba Island). In addition, two of the species, N. aguirrei sp. nov. and N. sona sp. nov., described from Isla Canales de Afuera, were found at different locations on the Soná Peninsula. Subject to additional collecting, we can say that the data suggest some speciation has occurred in isolation on Isla Canales de Afuera. We would also note that other more common microcaddisfly species found in different locations throughout western Panama are also present on that island (e.g. Neotrichia armata Botosaneanu, N. carlsoni Harris & Armitage and N. xicana (Mosely)).
The addition of five more species of the genus Neotrichia continues a remarkable story which seemingly has no end. Starting with three recorded species when we started our research in Panama in 2014, we have added 42 new species and new country records. Now that total is 50, including the five new species described herein.
Similarly, the number of caddisfly taxa in Panama shows no signs of slowing down. The 257 species recorded when we began our work increased to 535, more than double the number, by 2024. With the publication of this manuscript, Panama now boasts 544 caddisfly species.
Two of the species (N. aguirrei sp. nov. and Metrichia riosi sp.nov.) were named for aquatic biologists at MUPADI in recognition of their selfless work to make the progress and successes, which AIRG has experienced, a reality. In several previous publications (
Finally, the results in this paper are also a significant contribution to our knowledge of caddisfly distributions in Panama. Before this, no adult taxa were recorded in the refereed literature from Coiba National Park and only a few species, based on AIRG work, from the Soná Peninsula. Considering that most areas in Panama have never been collected, we anticipate that much more remains to be learned about the fauna and distribution of caddisflies in Panama.
We are indebted to the Rector and Vice-Rector of the Universidad Autónoma de Chiriquí, David, Panamá for encouraging our research and certifying our various projects. Our ongoing thanks to the Ministry of the Environment (MiAmbiente) in Panama for providing collecting permits, import/export permits and other forms of support. They also served as the primary manager of the Sustainable Production System and Biodiversity Conservation Project (PSPSCB) project, funded by the World Bank, with which the first two authors were involved and which provided us with useful specimens of Ochrotrichia martinez n.sp. from several of Panama’s national parks and protected areas. We appreciate the organisational and logistical support by the Gorgas Institute and COZEM during the PSPSCB project, as well as their field crews for collecting those specimens. We thank Kayla Sanchez Castillo, Marietta Landis, Tomás A. Ríos González, Yusseff P. Aguirre, Aydeé Cornejo, Edgar Pérez, Eric Álvarez and Carlos Nieto for their efforts in field collecting. We are very appreciative of the editing skills performed by Tatiana I. Arefina-Armitage on the manuscript. Funding for this project and report was made possible thanks to support for Dr Brian J. Armitage by the Sistema Nacional de Investigación (SNI) of Secretaría Nacional de Ciencia, Tecnología e Innovación (SENACYT), Republic of Panama.
The authors have declared that no competing interests exist.
No ethical statement was reported.
We acknowledge support for this work and report made to the second author by the Sistema Nacional de Investigación (SNI) of SENACYT, Republic of Panama.
Conceptualisation: BJA, VR. Data curation: BJA, SCH. Formal analysis: BJA. SCH, VR. Writing and editing (BJA, SCH). Funding acquisition: BJA.
Brian J. Armitage https://orcid.org/0000-0003-3182-1533
Steven C. Harris https://orcid.org/0000-0002-6432-7462
Viterbo Rodriguez https://orcid.org/0000-0003-1592-4479
All of the data that support the findings of this study are available in the main text.