Description of a new species of Trichophoromyia (Diptera: Psychodidae: Phlebotominae) from the Amazonian region of Maranhão, Brazil

Descrição de uma nova espécie de Trichophoromyia (Diptera: Psychodidae: Phlebotominae) da Amazônia Maranhense, Brasil

Abstract

Sand flies include vector species of protozoa of the genus Leishmania Ross, 1903. The genus Trichophoromyia Barretto, 1962 has a wide distribution in the Amazon biome, where several species occur in both preserved and anthropized environments, some of which have been incriminated or are suspected vectors of cutaneous leishmaniasis. In this study, we describe a new Trichophoromyia species, from males collected in the Gurupi Biological Reserve, located in the municipality of Centro Novo do Maranhão, state of Maranhão, Brazil. The description is based on morphological and morphometric characters of three male specimens captured with CDC-type light traps. The new species is distinguished from the other species of the genus mainly by having a set of two well-calibrated setae, two mid-caliber setae, and fine setae in the baso-median region of the gonocoxite, in addition to characteristics of the paramere and the aedeagal ducts. With the description of this new taxon, the number of valid species of the genus increases to 49, expanding the taxonomic knowledge about it.

Keywords:
Phlebotominae; forest; cutaneous leishmaniasis; biological reserve; taxonomy

Resumo

Os flebotomíneos incluem espécies vetoras de protozoários do gênero Leishmania Ross, 1903. O gênero Trichophoromyia Barretto, 1962 apresenta ampla distribuição no bioma Amazônia, onde várias de suas espécies ocorrem em ambientes florestais e antropizados, sendo algumas incriminadas ou suspeitas de transmitir agentes etiológicos das leishmanioses tegumentares. Neste estudo, descrevemos uma nova espécie de Trichophoromyia, com base em machos coletados na Reserva Biológica do Gurupi, localizada no município de Centro Novo do Maranhão, estado do Maranhão, Brasil. A descrição fundamenta-se em caracteres morfológicos e morfométricos de três espécimes machos capturados com armadilhas luminosas do tipo CDC. A nova espécie distingue-se das demais espécies do gênero principalmente por apresentar, na região basomediana do gonocoxito, um conjunto formado por duas cerdas robustas, duas cerdas de calibre intermediário e cerdas finas, além de características particulares do parâmero e dos ductos edeagais. Com a descrição desse novo táxon, o número de espécies válidas do gênero passa a 49, ampliando o conhecimento taxonômico sobre esse grupo.

Palavras-chave:
Phlebotominae; floresta; leishmaniose tegumentar; reserva biológica; taxonomia

1. Introduction

Sand flies (Diptera, Psychodidae, Phlebotominae) are insects of medical and veterinary importance, as they are vectors of bacteria, viruses, and protozoa of the genus Leishmania Ross, 1903 (Akhoundi et al., 2016). There are 1,067 described taxa in the world, of which 33 are fossils (Galati et al., 2025). In the Western Hemisphere, sand flies comprise 556 species, including 539 extant and 17 fossil species, assigned to 25 genera (Galati et al., 2025; Rodrigues et al., 2026a).

Among these, 97 species/subspecies have been recorded for the state of Maranhão, making it one of the most diverse regions for sand flies in Brazil (Rebêlo et al., 2010; Aguiar and Vieira, 2018; Rodrigues and Galati, 2024; Galati, 2026). The western portion of Maranhão lies within the Amazon biome, one of the world's major biodiversity hotspots. This environmental heterogeneity is reflected in its rich phlebotomine fauna, with the highest sand fly species richness in Brazil occurring in Amazonian states (Aguiar and Vieira, 2018).

The genus Trichophoromyia Barretto, 1962 was originally proposed as a subgenus of Lutzomyia França, 1924, and later elevated to the genus level by Galati (1995). According to Galati, Trichophoromyia belongs to the subtribe Psychodopygina Galati, 1995, alongside six other genera: Psathyromyia Barretto, 1962, Viannamyia Mangabeira 1941, Martinsmyia Galati, 1995, Bichromomyia Galati 1995, Psychodopygus Mangabeira, 1941, and Nyssomyia Barretto, 1962 constitute the subtribe Psychodopygina Galati, 1995. Recently, the generic composition of this subtribe was expanded with the description of two new genera: Shawmyia Rodrigues & Galati, 2026 and Reburramyia Rodrigues & Galati, 2026 (Rodrigues et al., 2026a, b), the later composed by a singles species previously classified as a Trichophoromyia. Additionally, Trichophoromyia was subdivided into the subgenera Trichophoromyia s. str. and Dilermandomyia Rodrigues & Galati, 2026 (Rodrigues et al., 2026a).

Most species of Trichophoromyia occur in the Amazon biome, inhabiting both preserved forests and human-modified environments (Aguiar and Vieira 2018). Females of the subgenus Dilermandomyia are highly isomorphic, making species-level identification difficult. Consequently, species delimitation and taxonomy rely primarily on male morphological characters.

The genus has also gained increasing epidemiological importance because Trichophoromyia ubiquitalis (Mangabeira, 1942) is the confirmed vector of Leishmania (Viannia) lainsoni Silveira, Shaw, Braga e Ishikawa, 1987, while additional species of Dilermandomyia have been implicated as suspected vectors of other agents of cutaneous leishmaniasis (Santos and Silveira 2020; Rodrigues et al., 2026a).

In this study, we describe a new species of Trichophoromayia (Dilermandomyia) based on male morphology. The specimens were collected in the Gurupi Biological Reserve (REBIO Gurupi), located in the municipality of Centro Novo do Maranhão, state of Maranhão, within the Eastern Amazon region of Brazil.

2. Methodology

2.1. Study area

The Gurupi Biological Reserve (REBIO-Gurupi) is located between coordinates 3°09’S to 4°08”S and 46°27”W to 46°58”W in the northwest region of the State of Maranhão, between the basins of the Gurupi and Pindaré Rivers. It is the only Federal Conservation Unit (CU) in this state covering an area of 271,197 ha (Brasil, 2000) (Figure 1). It is of great importance for the conservation of this forest, located in the Belém Endemism Center, and is included in the list of priority conservation areas (Almeida and Vieira, 2010).

Figure 1
Area of the Gurupi Biological Reserve highlighting the collection point of sandflies (yellow dot) captured between May and June 2022, Maranhão, Brazil.

This CU borders the Awá and Caru indigenous lands to the east, and the Gurupi River, to the north (IBAMA, 1999). The region has a tropical rainy climate, characterized by two well-defined seasons: a rainy season (from December to May) and a dry season (from June to November). The altitude varies from 15 to 315 m. The average annual rainfall varies from 1,750 to 2,000 mm, and the temperature is between 24 and 26 °C (Birdlife International, 2016).

The area has transitional vegetation (Martins and Oliveira 2011), with a predominance of Submontane Dense Ombrophilous Forest, representing 3.10% of the region where the REBIO-Gurupi is located, and has 17.08% of secondary vegetation (IBGE, 2019). Its floristic richness indicates the peculiarity of this forest remnant, which serves as an ecological space for several species of wild fauna, such as mammals, reptiles, and birds, with species threatened with extinction or with reduced geographical distribution (Hessel and Lisboa, 2015).

The research group has the Permanent License for the Collection of Zoological Material, issued by the Biodiversity Authorization and Information System (SISBIO, process No. 97762-2).

2.2. Morphology

The description of the new species is based on morphological and morphometric characters of three males captured in the REBIO-Gurupi, located in Centro Novo do Maranhão, Maranhão, Brazil. CDC-type light traps were used and operated from 18:00 to 06:00 hours. The holotype was collected on 2022-V-29 (03°30’35.1”S 046°49’44.6”W), one paratype on 2022-V-30 (03°30’55.8”S 046°49’33.6”W), and the other paratype on 2022-VI-03 (03°30’55.8”S 046°49’33.6”W). Drawings were made using a camera lucida attached to the Leitz Dialux 22 optical microscope, and digitized using Adobe Illustrator version 26.0. All measurements are given in micrometers (μm). The microscope measurements were calibrated as follows: Calibration factors were 6.3x = 18.5 μm; objective 16x = 7.3 μm; objective 25x = 4.65 μm; 40x objective = 2.9 μm. Measurements outside parentheses refer to the holotype, whereas those in parentheses indicate the range observed in the paratypes. The sample size (n) refers to the number of paratypes examined.

Descriptions of the species T. (D.) brachipyga (Mangabeira 1942), T. (D.) gibba (Young and Arias, 1994) and T. (D.) viannamartinsi (Sherlock and Guitton, 1970) were consulted. The specimens of the new species were compared with photos of the paratype males of T. (D.) brachipyga and T. (D.) gibba deposited in the Coleção Entomológica de Referência da Faculdade de Saúde Pública da USP and in the Coleção de Flebotomíneos (COLFLEB), Instituto René Rachou, Fiocruz Minas, Belo Horizonte, Minas Gerais, Brazil, respectively. To classify Phlebotominae, we adopted the classification proposed by Galati (2018). The nomenclature of characters and terminology are in accordance with Sábio et al. (2014) and Galati et al. (2017).

3. Results

Trichophoromyia (Dilermandomyia) Gurupi Rodrigues, Chaves-Júnior, Galati & Pinheiro sp. nov.

(Figures 2 -4)

Figure 2
Male holotype of Trichophoromyia gurupi sp. n. (A) Head; (B) Flagellomere I; (C) Flagellomere II; (D) Flagellomere III; (E) Palpomere I; (F) Palpo II; (G) Palpo III; (H) Palpo IV. (Bars: 100 μm).

Figure 3
Male holotype of Trichophoromyia gurupi sp. n. (A) Wing; (B) Spermatic pump and edeagal ducts (paratype). (Bar: 100 μm).

Figure 4
Male holotype of Trichophoromyia gurupi sp. n. (A) Genitalia; (B) Parameres in different positions; (C) Aedeagal ducts and parameral sheath. (Bars: 100 μm).

3.1. Classification

• Phylum Arthropoda von Siebold, 1848.

• Subphylum Hexapoda Latreille, 1825.

• Class Insecta Linnaeus, 1758.

• Order Diptera Linnaeus, 1758.

• Suborder Psychodomorpha Hennig, 1968.

• Family Psychodidae Newman, 1834.

• Subfamily Phlebotominae Rondani and Berté, 1840, in Rondani, 1840

• Genus Trichophoromyia Barretto, 1962.

• Subgenera Dilermandomyia Rodrigues & Galati, 2026

  1. T

     (D) gurupi Rodrigues, Chaves-Júnior, Galati & Pinheiro sp. nov.

3.2. Diagnosis

The male of T. (D.) gurupi sp. nov can be differentiated from the other species of the genus Trichophoromyia, because it has a cluster in the gonocoxite with two very large setae, two medium-caliber setae, and other thin ones (Figure 4A).

3.3. Male holotype

Insect of predominantly brown color, with part of the pleurae pale. Dark brown pronotum, mesonotum, and metanotum. Katepisternum and Katepimeron brown. Paratergite, light brown upper anepisternum.

Head (Figure 2A): length 363 μm (340 – 363, n = 2), width 305 (336, n = 1); head length/width ratio 1.19 (1.08, n = 1). Eyes, length 206 (189, n = 2), width 101 (104 – 110, n = 2). Incomplete interocular suture, interocular distance 110 (116, n = 1). Pharynx with stretch marks without teeth. Cibarium without teeth and an incomplete sclerotized arch. Clypeus length 113 (110, n = 2), width 64 (64 – 70, n = 2), labrum-epipharynx length 194 (191 – 203). Holotype antenna with only the first three flagellomeres intact, the others are deformed (paratypes without antennae). Antenna: FI: 217 (Figure 2B), FII: 113 (Figure 2C), FIII: 113 (Figure 2D). Pedunculated ascoids with very short posterior prolongation; long anterior extension in IF, exceeds the FII basis; external ascoid more basal than the internal ascoid. Presence of preapical papilla in FI and FII (Figures 22C), absent in FIII (Figure 2D). Palpus, length: PI 35 (35, n = 2), PII 93 (99 – 101, n = 2), PIII 119, PIV 46, PV lost. Palpal formula: 1.4.2.3. Newstead’s sensillas present in the second palpal segment (Figure 2F) and dispersed in the third palpal segment (Figure 2G). Labial suture forming a fork (Figure 2A).

Cervix – Absence of ventro-cervical sensillas. Presence of two sensilla in the cervical sclerites

Thorax: 547 (526 – 576, n = 2) long, mesonotum 489 (475 – 519, n = 2) long. Presence of 2 proepimeral setae (2-3, n = 2); 12 maxillary anepisternal setae (10, n = 2). Absence of setae in the anterior region of the Katepisternum. Wing (Figure 3A): length 2,054, width 540; Length/width ratio 3.80. Veins: R₅ 1.258; Alpha 504 (526, n = 1), Beta 261 (264, n = 1), Gamma 261 (255, n = 1), Delta 328 (319, n = 1), Pi 88 (117, n = 1). The legs: only the femora were preserved for the holotype and paratypes, in addition to an anterior and middle femur, as the other segments were lost in the clarification process. Coxa length (anterior, median, and posterior, respectively): Coxa 335 (348, n = 2), 335 (335 – 353, n = 2), 353 (345 – 349 n = 2); Femur 795 (832, n = 1), 758 (777, n = 1).

Abdomen: the first three abdominal segments were removed for DNA extraction. However, DNA extraction was unsuccessful in two individuals, preventing the acquisition of DNA of sufficient quality for COI gene-based DNA barcoding. Absence of tergal papillae. Genitalia (Figure 4A): Gonocoxite (Figure 5A); length 446 (418 – 428 n = 2), width 228 (190 – 205, n = 2), presence of a basal tuft of setae, two of which are very large and with a hook-shaped apex, two medium-caliber setae, and fine setae. Gonostyle 293 (260 – 265, n = 2), with four spines and absence of the preapical spiniform bristle. Arrangement of the spine: an apical one; the superior external in a subapical position and the inferior external implanted after the median region, so that it is closer to the superior than the internal, the latter implanted in the first basal third of the gonostyle. Paramere (Figure 4B); dorsal margin length 148 (148 – 154, n = 2), ventral margin 142 (142 – 145, n = 2); presence of median lobe covered by thin setae; two setae with hooked apex in the post-lobe region; apex of the rectangular paramere, covered by fine setae. Triangular parameral sheath, it was not possible to measure the margins of the parameral sheath by the position in which it is. Epandrial lobe: 532 (525 – 554, n = 2) long, 38 (35, n = 2). Sperm pump absent in the holotype, measurements are from the paratype: lenght (148, n =1). Ejaculatory apodeme: lenght (113, n = 1). Incomplete aedeagal ducts in all specimens; apex of the tapered aedeagal ducts (Figure 4C).

Figure 5
Male holotype of Trichophoromyia gurupi sp. n. (A) Gonocoxite isolated to represent the tuft of bristles (Bars: 100 μm).

3.4. Material examined

The holotype and paratypes were collected in the Gurupi Biological Reserve (REBIO-Gurupi), in the municipality of Centro Novo do Maranhão, Maranhão State, Brazil, using CDC light traps. The holotype collected on: V/29/2022 (03°30’35.1’’S 046°49’44.6’’W), and the two paratypes at the same locality (03°30’55.8’’S 046°49’33.6’’W), but on different days paratype 1: V/30/2022 and paratype 2: VI/03/2022. The holotype and two paratypes are deposited in the Coleção de Flebotomíneos (COLFLEB), Instituto René Rachou, Fiocruz Minas, Belo Horizonte, Minas Gerais, Brazil. The specimens were examined and approved through the collection's critical analysis process and are registered under accession numbers 92580 (holotype), 92581, and 92582 (paratypes).

3.5. Etymology

The specific epithet gurupi is a noun in apposition, derived from the toponym Gurupi, in reference to the Gurupi Biological Reserve, located in the municipality of Centro Novo do Maranhão, western region of the state of Maranhão, Brazil, where the type material of the new species was collected.

Geographical Distribution: BRAZIL. Maranhão: REBIO do Gurupi, Centro Novo do Maranhão.

4. Discussion

4.1. Taxonomic discussion

Due to the high morphological similarity among females of the genus Trichophoromyia, many species are known exclusively from males. Morphologically, males are characterized by terminalia with a length equal to or greater than that of the thorax, and a gonostyle bearing short spines, with the apical spine usually shorter than the gonostyle itself (Galati, 2026). Morphological identification of females is possible for less than 10% of its species (Galati, 2018).

T. (D.) gurupi sp. nov. possesses aedeagal ducts four times as long as the sperm pump. This characteristic place the species within the subgenus Dilermandomyia, distinguishing it from species of the subgenus Trichophoromyia, in which this ratio is equal to or less than three times (Rodrigues et al., 2026a).

T. (D.) gurupi sp. nov. has a voluminous gonocoxite (length equal to or less than 2.2 times its width) with a basal-median tuft containing two setae that are thicker and more sclerotized than the rest. Additionally, the ventral articulation area between the gonostyle and gonocoxite is well developed and strongly striated. Based on these characteristics, T. (D.) gurupi sp. nov. closely resembles T. (D.) bettini (Le Pont and Desjeux, 1987), T. (D.) brachipyga (Mangabeira, 1942), T. (D.) eurypyga (Martins, Falcão & Silva, 1963), T. (D.) gibba, T. (D.) jariensis Cavalcante, Rodrigues & Galati, 2024, T. (D.) lopesi (Damasceno, Causey & Arouck, 1945), and T. (D.) viannamartinsi.

The new species differs from T. (D.) lopesi because the latter possesses a trilobed paramere. It differs from T. (D.) jariensis, which exhibits a tuft of approximately 20 setae on the basal inner margin of the gonocoxite. It can be distinguished from T. (D.) bettini because the latter lacks a tubercle with thin setae at the base of the dorsal lobe of the paramere, features a nearly straight posterior region relative to this lobe, and has a rounded paramere apex. Conversely, the presence of a tubercle with thin setae at the base of the dorsal lobe, a pronounced concavity posterior to this lobe, and a preapical area of the paramere facing the gonocoxite aligns with T. (D.) gurupi sp. nov. with T. (D.) brachipyga, T. (D.) eurypyga, T. (D.) gibba, and T. (D.) viannamartinsi.

Among these, T. (D.) gibba differs by having a row of setae on the dorsal margin of the paramere located between the posterior region of the median lobe and the apex, whereas this area is glabrous in T. (D.) gurupi sp. nov., T. (D.) brachipyga, T. (D.) eurypyga, and T. (D.) viannamartinsi. Furthermore, T. (D.) eurypyga is distinguished from these species by a basal-median tuft on the gonocoxite composed of about 30 setae of uniform caliber. In T. (D.) brachipyga, this tuft contains five setae that are much thicker and more sclerotized than the others, whereas T. (D.) gurupi sp. nov. and T. (D.) viannamartinsi possess only two such setae.

In the new species, in addition to the two large-caliber, highly sclerotized setae, there are two other smaller-caliber sclerotized setae, as well as two groups of thin setae positioned above and below the sclerotized ones. In contrast, T. (D.) viannamartinsi possesses only the two large-caliber sclerotized setae and a single continuous group of approximately 20 thin setae located more ventrally than the sclerotized ones (Galati, 2026).

The geographical distribution of the species morphologically related to T. (D.) gurupi sp. nov. is characterized as follows: T. (D.) bettini [Venezuela]; T. (D.) brachipyga [French Guiana, Brazil (Roraima, Amapá, Amazonas, Pará, Acre, Rondônia)], T. (D.) eurypyga [(Venezuela, Brazil (Roraima, Amapá, Amazonas, Pará, Rondônia)], T. (D.) gibba [Brazil (Amazonas)], T. (D.) jariensis [Brazil (Amapá)], T. (D.) lopesi [Brazil (Amazonas)], and T (D.) viannamartinsi [Brazil (Maranhão, Pernambuco and Bahia)] (Galati, 2026). The distribution of these morphologically related species reveals a predominantly Amazonian pattern, extending into transitional areas of northern and northeastern South America.

5. Conclusion

The description of T. (D.) gurupi sp. nov. not only expands the known species diversity of the genus but also represents an original contribution to the taxonomy of Trichophoromyia by providing a new morphological characterization from the Amazonian region of Maranhão, Brazil. The discovery of this new species highlights the still underestimated taxonomic diversity of sand flies in the region and underscores the importance of continuing entomological surveys in in poorly studied areas, which remain essential for deepening our understanding of taxonomy, biogeography, and evolutionary relationships within Trichophoromyia. Currently, the genus comprises 49 recognized species, 28 of which have been recorded in Brazil. The description of T. (D.) gurupi sp. nov. therefore represents an important contribution to our knowledge of the Brazilian fauna and provides new morphological information that can support future comparative and integrative taxonomic studies of the genus. Furthermore, this discovery underscores the importance of continued sampling in the Amazon biome, where additional undescribed species may still occur, particularly in areas that remain poorly explored from a taxonomic and entomological perspectives.

Acknowledgements

The authors are grateful to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and the Fundação de Amparo à Pesquisa e ao Desenvolvimento Científico e Tecnológico do Maranhão (FAPEMA) for their financial support through the Programa de Apoio a Projetos de Pesquisa para a Capacitação e Formação de Recursos Humanos em Taxonomia Biológica (PROTAX, Edital nº 24/2024) and the Chamada Pública CNPq/CONFAP nº 34/2023 – Expedições Científicas – Iniciativa Amazônia +10. The authors also thank the Universidade Estadual do Maranhão (UEMA) and the Laboratório de Entomologia Médica (LABEM/UEMA) for providing the infrastructure necessary to conduct this research; the Universidade de São Paulo (USP), especially the Laboratório de Entomologia em Saúde Pública (LESP/USP); and the Universidade Federal do Paraná (UFPR), particularly the Laboratório de Parasitologia Molecular, for their technical and scientific support and valuable collaboration in the comparative analyses. The authors also express their sincere gratitude to Prof. Dr. Andrey José de Andrade for his invaluable assistance and guidance throughout the process of describing the new species.

Data Availability Statement

The entire data set that supports the results of this study was published in the article itself.

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e-mail: [e-mail]

Editor:

Takako Matsumura Tundisi

Publication Dates

• Publication in this collection
  09 Oct 2026
• Date of issue
  2026

History

• Received
  10 July 2026
• Accepted
  05 Sept 2026