Research Article |
Corresponding author: Pedro Bonfá-Neto ( bonfa.pn@gmail.com ) Academic editor: Steffen Pauls
© 2024 Pedro Bonfá-Neto, Frederico Falcão Salles, Albane Vilarino.
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:
Bonfá-Neto P, Salles F, Vilarino A (2024) New species of the long-horned caddisfly Oecetis McLachlan, 1877 (Trichoptera: Leptoceridae) from the Atlantic Forest, Brazil and their evolutionary relationship. Arthropod Systematics & Phylogeny 82: 551-566. https://doi.org/10.3897/asp.82.e114286
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Asymmetrical genitalia are reported from major Trichoptera subgroups and evolved multiple times independently. In Oecetis, it is a characteristic of the insconpicua group. However, certain species in other species groups also evolved an asymmetrical spiny projection on the phallotheca. Here, two new species with an asymmetric projection are described in the falicia group from the Brazilian Atlantic Forest, Espírito Santo state. Their phylogenetic relationships were investigated through a Bayesian analysis combining COI and morphological data. Additionally, we provide new records of Oecetis connata, O. inconspicua and O. paranensis from the Espírito Santo state, and O. connata and O. flinti from Minas Gerais state. Oecetis capixaba sp. nov. is placed with low support as the sister species of O. acanthostema, both presenting stout spine-like setae on the inner surface of the inferior appendage; the new species is diagnosed by the long spine-like setae on the inferior appendage, the narrow dorsolateral process of segment IX, and the phallic apparatus without apical projections. Oecetis ruschii sp. nov. is placed as a sister group of the clade including O. facilia and O. furcata, both presenting forked dorsolateral processes of segment IX; the new species is diagnosed by the dorsolateral process of segment IX with a lateral branching and the apex of inferior appendage wide and triangular. While the phylogenetic results should be considered preliminary and interpreted with caution, they indicate that the asymmetric projection evolved multiple times convergently in the avara, punctata, and falicia groups. The asymmetric genitalia in the falicia group seems to have evolved only in males since no correspondent asymmetry is described for females. The function of the asymmetrical projection remains unknown.
Aquatic insects, Asymmetric genitalia, Bayesian inference, Diversity, Neotropical
Oecetis McLachlan, 1877 is the only member of the tribe Oecetini Silfvenius, 1905 of the family Leptoceridae Leach, 1815. The genus is distributed worldwide, with their immatures inhabiting major rivers and lotic environments where they usually are very common and abundant (
The relationship among the long-horned caddisflies (Leptoceridae) was investigated by
Several species groups have been proposed for Oecetis (
The phylogenetic relationship of the Neotropical Oecetis was investigated through morphological characters by
Here we describe and illustrate two additional falicia group species from the Atlantic Forest, Brazil. Both new species present a conspicuous asymmetrical process on the phallic apparatus. Asymmetrical genitalia have been observed in all major Trichoptera lineages and have evolved convergently numerous times (
The specimens were collected with various types of light traps (UV pan traps, Pennsylvania traps, and light over a white cloth) or Malaise traps as indicated in the material analyzed sections. The morphological terminology follows
For the phylogenetic placement of the new species we modified the morphological matrix of
List of morphological characters and states, for Oecetis and related taxa. (Modified from
Character list and states: | |
---|---|
0 | Forewing, M vein: (0) Branched into M 1+2 and M 3+4 ; (1) Unbranched. |
1 | Middle leg femur, row of spines on the inner surface: (0) Absent; (1) Present. |
2 | Middle leg femur spines, coverage area: (0) The whole podomere; (1) Half of the podomere length. |
3 | Hind leg tibia, row of spines on the inner surface: (0) Absent; (1) Present. |
4 | Hind leg tibia spines, coverage area: (0) The whole podomere; (1) Half of the podomere length. |
5 | Foreleg tibia, apical spur: (0) Absent; (1) Present. |
6 | Forewing, fork V: (0) Rooted; (1) Sessile; (2) Petiolate. |
7 | Forewing,sectoral crossvein (r 2+3 –r 4+5) alignment to the r–m crossvein: (0) Aligned; (1) not-aligned. |
8 | Hind wing, Rs sector on the hind wing: (0) Absent; (1) Present. |
9 | Hind wing, fork I: (0) Absent; (1) Present. |
10 | Forewing, setae fringe on the inner surface: (0) Absent; (1) Present. |
11 | Forewing, end of Sc vein: (0) Vestigial; (1) Whole. |
12 | Forewing, black spots at the end of veins, forks and junctions on the membrane: (0) Absent; (1)Present. |
13 | Forewing, dark bands on the membrane around the cord: (0) Absent; (1) Present. |
14 | Forewing, apex: (0) Rounded; (1) Acuminated. |
15 | Hind wing, position of r–m crossvein related to the fork between M 1+2 and M 3+4: (0) Rooted; (1) Sessile; (2) Petiolate. |
16 | Forewing, fork I in relation to the discoidal cell crossvein: (0) Rooted; (1) Sessile; (2) Petiolate. |
17 | Forewing, depth of the fork I: (0) Shallow; (1) Deep. |
18 | Hind wing, anal region: (0) Wide (as in Ceraclea and Athripsodes); (1) Narrow (as in Brachysetodes and Setodes). |
19 | Tergum IX related to the sternum IX, length: (0) Longer than sternum IX; (1) same length of sternum IX; (2) shorter than sternum IX. |
20 | Terga V to VIII honeycomb texture: (0) Absent; (1) Present. |
21 | Tergum IX and X, acrotergite: (0) Absent; (1) Present. |
22 | Tergum IX and X acrotergite, number: (0) One; (1) Two. |
23 | Segment IX, dorsolateral processes: (0) Absent; (1) Present. |
24 | Segment IX, dorsolateral processes curvature: (0) Straight; (1) Bent ventrally. |
25 | Segment IX, dorsolateral processes relative length: (0) As long as the preanal appendages; (1) Much longer than the preanal appendages. |
26 | Segment IX, dorsolateral processes shape: (0) Thread-like; (1) Forked. |
27 | Preanal appendage fusion: (0) Completely fused to each other; (1) Not fused to each other; (2) Partially fused to each other. |
28 | Preanal appendages, shape in dorsal view: (0) Ovoid; (1) Digitate. |
29 | Tergum X, dorsal lobe: (0) Absent; (1) Present. |
30 | Tergum X, dorsal lobe shape: (0) Cylindrical throughout length; (1) Cylindrical with globose apex; (2) Flat; (3) Saddle-shaped. |
31 | Tergum X, median incision: (0) Absent (undivided tergum X); (1) Present (divided tergum X). |
32 | Tergum X, lobes of divided tergum, shape in dorsal view: (0) Rod-like; (1) Broad at base and tapering distally. |
33 | Tergum X, apex: (0) Truncate; (1) Acuminate; (2) Irregular shape; (3) Round. |
34 | Tergum X median incision shape: (0) V-shaped shallow; (1) V-shaped deep; (2) U-shaped. |
35 | Inferior appendages, length: (0) Short; (1) Long. |
36 | Inferior appendages, dorsal lobe: (0) Absent; (1) Present. |
37 | Inferior appendages, dorsal lobe shape: (0) Quadrate; (1) Ovoid; (2) Digitate; (3) Triangular; (4) L-shaped. |
38 | Inferior appendages, dorsal lobe insertion: (0) Inner margin; (1) Laterally. |
39 | Inferior appendages, dorsal lobe orientation angle: (0) Projecting upward (90 degrees to the distal portion); (1) Projecting distally (less than 90 degrees to the distal portion). |
40 | Inferior appendages, ventral lobe: (0) Absent; (1) Present. |
41 | Inferior appendages, ventral lobe shape: (0) Quadrate; (1) Digitate; (2) Triangular; (3) acute |
42 | Inferior appendages, ventral lobe size: (0) Small (less than 1/4 length of distal portion); (1) big (more than 1/4 length of distal lobe). |
43 | Inferior appendages, general shape: (0) Cylindrical with apex rounded; (1) Tapering distally with apex acute; (2) Short and “fist-like”; (3) Cylindrical proximally, enlarged distally. |
44 | Inferior appendages, when short, fist-like, distal lobe shape: (0) ovoid, ear-like with smooth and rounded edges; (1) quadrate with thick setae on apex; (2) Cylindrical and stout. |
45 | Inferior appendages, distal lobe, apical incision: (0) Absent; (1) Present. |
46 | Inferior appendages, distal lobe apical incision, shape in lateral view: (0) V-shaped; (1) L-shaped. |
47 | Inferior appendages, inner lobe: (0) Absent; (1) Present. |
48 | Segment IX, posterolateral margin, setae: (0) Absent; (1) Present. |
49 | Inferior appendages, thick spine-like setae: (0) Absent; (1) Present. |
50 | Phallic apparatus, symmetry in dorsal view: (0) Symmetrical; (1) Asymmetrical. |
51 | Phallic apparatus, length: (0) Short; (1) Elongated. |
52 | Phallic apparatus, when short, shape: (0) Comma-shaped, strongly bent ventrally; (1) Round, inflated; (2) Cylindrical, slightly bent ventrally. |
53 | Phallic apparatus, when short, ventrodistal lamellate process: (0) Absent; (1) Present. |
54 | Phallic apparatus, endotheca, size: (0) Small (less than 1/3 the phallus volume); (1) Large (more than 1/3 the phallus volume). |
55 | Phallic apparatus, endotheca, number of lobules: (0) One; (1) Two. |
56 | Phallic apparatus, additional sclerotized structure in phallic apparatus (other than phallic spine or phallotremal sclerite): (0) Absent; (1) Present. |
57 | Phallic apparatus, phallic spines: (0) Absent; (1) Present. |
58 | Phallic apparatus, phallic spines, number: (0) One; (1) Two; (2) Three. |
59 | Phallic apparatus, phallic spines, shape: (0) Straight; (1) Curved. |
60 | Phallic apparatus, phallotremal sclerite: (0) Absent; (1) Present. |
61 | Phallic apparatus, phallotremal sclerite, number: (0) One; (1) Two. |
62 | Phallic apparatus, phallotheca, asymmetric spine projection: (0) Absent; (1) Present. |
63 | Phallic apparatus, when short, shape: (0) Comma-shaped, strongly bent ventrally; (1) Round, inflated; (2) Cylindrical, slightly bent ventrally. |
Sequences of COI barcode used in the phylogenetic inference, and respective BOLD accession number.
Species | BOLD accession number |
Athripsodes bergensis | SATRI002-13 |
Brachysetodes major | UMNEB140-08 |
Leptocerus americanus | AVMTT037-09 |
Mystacides interjectus | ABCAD059-08 |
Nectopsyche argentata | UMNEB177-08 |
Nectopsyche punctata | GBMIN35590-13 |
Setodes incertus | BKCAD041-08 |
Oecetis akimi | GBMNB60844-20 |
Oecetis amazonica | GBMNB60805-20 |
Oecetis arizonica | OFCAD375-08 |
Oecetis avara | HIEPT040-09 |
Oecetis connata | GBMNB60798-20 |
Oecetis excisa | GBMNB60809-20 |
Oecetis iguazu | KKUMN207-10 |
Oecetis inconspicua | GBMNB60806-20 |
Oecetis inscripta | UMNEA325-08 |
Oecetis knutsoni | UMNEB251-08 |
Oecetis lacustris | BARCO038-14 |
Oecetis marquesi | TRHGO332-10 |
Oecetis metlacensis | UMNEA311-08 |
Oecetis nigropunctata | RUSST215-12 |
Oecetis ochracea | UMNEB664-08 |
Oecetis paranensis | KKUMN211-10 |
Oecetis pechana | AUCAD016-09 |
Oecetis pratti | OFTRI403-10 |
Oecetis prolongata | KKUMN212-10 |
Oecetis pseudoinconspicua | UMNEA332-08 |
Oecetis punctata | UMNEA333-08 |
Oecetis punctipennis | GBMNB60810-20 |
Oecetis testacea | CAUTR083-09 |
The phylogeny was based on a probabilistic framework through Bayesian inference as implemented in MrBayes 3.2.7 (
Morphological partitions used for the Bayesian inferences. The partitions were established based on its levels of homoplasy obtained from the adjusted homoplasy of a cladistic analysis under implied weight. Individual values were combined into more inclusive classes.
Partition | Adjusted homoplasy | Characters |
1 | 0 | 0, 8, 23, 25, 26, 38, 44, 46, 47, 52 |
2 | 0.1 | 9, 24, 27, 42, 55, 19, 20, 22, 58 |
3 | 0.2 | 3, 17, 56, 59, 15, 37, 39, 41, 45, 49, 62, 2, 40, 50, 51 |
4 | 0.3 | 35, 53, 12, 28, 32, 48, 54, 57, 21, 29, 30, 36 |
5 | 0.4 | 7, 34, 6, 31, 43, 5 |
6 | 0.5 | 14, 60, 1, 16, 13, 33 |
7 | — | 4, 10, 11, 18, 61 |
The COI sequences available in the BOLD website (Barcode of life Database) were included to provide additional evidence about taxa relationship. We included in the phylogenetic analyses COI sequence fragments for 30 species (Table
The analyses were performed through the CIPRES gateway (
The distributional map was created in QGIS Firenze ver. 3.28 software, using shapefile vector layers from the Instituto Brasileiro de Geografia e Estatística (IBGE) and the
The maximum credibility Bayesian tree obtained from the morphological characters and COI is presented in Figure
Maximum credibility Bayesian tree obtained from 63 morphological characters and 30 COI sequences coded to Oecetis and related taxa (all compatible groups shown). Morphological characters in DELTRAN optimization are displayed along the branches. Black symbols indicate unique character changes. Thick lines indicate lineages with asymmetrical spine projection on the phallotheca. Posterior probability support values are displayed near the node branches, values greater than 50% (majority consensus) are highlighted in orange, strongly supported clades (>90%) are shown in red. Taxa with included COI data are highlighted with *. Non-Neotropical Oecetis species distribution are indicated with the following abbreviations: AF Afrotropical, AU Austral, OR Oriental, PA Palearctic, NA Nearctic.
The avara group is indicated as monophyletic with very low support (PP = 16). The punctata group is placed in a distinct clade (PP = 61) and not within the avara group as in
The cladistic definition of the species groups according with the results are based on the following morphological characters: avara group: endotheca small (54:0), forewing fork I sessile (16:1), preanal appendage digitate (28:1) (PP = 16); inconspicua group: phallic apparatus asymmetrical (50:1), phallotremal sclerite curved (59:1), phallic apparatus round, inflated (52:1) (PP = 93); pratti group: mid leg femur with row of spines (1:1), forewing sectorial crossvein aligned (7:0), forewing apex acuminate (14:1), segment IX posterolateral margin setae absent (48:0) (PP = 10); punctata group: mid leg femur with row of spines (1:1), forewing fork I sessile (16:1), tergum X with irregular shape (33:2), inferior appendage ventral lobe small (42:0), inferior appendage quadrate, with thick setae (44:1) (PP = 61); punctipennis group: forewing fork V sessile (6:1), forewing apex acuminate (14:1), endotheca small (54:0) (PP = 98); falicia group: presence of dorsolateral process on the segment IX (23:1), tergum IX shorter than sternum IX (19:2), inferior appendage dorsal lobe triangular (37:3), and phallic apparatus elongate (51:1) (PP = 77).
Focusing on the falicia group (Fig.
Bayesian tree showing the Oecetis falicia group. Morphological characters in DELTRAN optimization are displayed along the branches; clades without supporting morphological characters were collapsed. Black symbols indicate unique character changes. Thick lines indicate lineages with asymmetrical spine projection on the phallotheca. Posterior probability support values are displayed near the node branches, values greater than 50% (majority consensus) are highlighted in orange, strongly supported clades (>90%) are shown in red. Taxa with included COI data are highlighted with *. Non-Neotropical Oecetis species distribution are indicated with the following abbreviations: AF Afrotropical, PA Palearctic, NA Nearctic. Illustrations of the male genitalia in lateral view based on original descriptions are depicted next to each species label, showing also for some species the lateral view of the phallic apparatus and the ventral view of the inferior appendage.
LEPTOCERIDAE Leach, 1815
Oecetis McLachlan, 1877
Holotype: BRAZIL • ♂; Espírito Santo, Santa Teresa, Augusto Ruschi biological reserve, Córrego Roda d’Água; 19°53′35.1″S, 40°32′39.7″W; 810 m a.s.l.; 24 Aug. – 30 Sep. 2017; Malaise trap; FF Salles, V Costa, P Bonfá Nt leg.;
This new species is similar to the other species of the falicia group with spine-like setae on the inner surface of the inferior appendage, being very similar to Oecetis achanthostema by both presenting a spiny process on the phalloteca. They can be differentiated by the dorsolateral process of segment IX in dorsal view being wide subapically in O. achanthostema, while it is tapered and overall narrow in the new species. The spine-like setae on the inferior appendage are conspicuously longer in the new species than in O. achanthostema. Additionally, the phallic apparatus of O. achanthostema has two digitate apical projections, which are absent in O. capixaba sp. nov.
Adult male: Forewing length 6.8 mm (n = 1). Head. Color pale light yellow (in alcohol). Antennae approximately 3x forewing length; scape stout, elongate; pedicel enlarged in width, narrower than scape, shorter than first flagellomeres; first flagellomere narrow, with same length as scape, other flagellomeres shorter than first. Maxillary palps pale light yellow, 5-segmented, segments subequal in length and width, densely covered with setae. Labial palps pale light yellow, apparently 4-segmented, first segment very small. Thorax. Pterothorax yellowish brown; forewing pale light yellow; dark bands over cord absent (Fig.
The specific epithet “capixaba” is a name originating from the Tupi language, meaning “farmland”, “a land clean for planting”, and it is currently used to designate people born in the state of Espírito Santo. Species named in apposition.
Holotype: BRAZIL • ♂; Espírito Santo, Santa Teresa, Augusto Ruschi biological reserve, Córrego Roda d’Água; 19°53′35.1″S, 40°32′39.7″W; 810 m a.s.l., 21–22 Oct. 2017; light pan trap; FF Salles, V Costa, P Bonfá Nt leg.;
This species is similar to Oecetis furcata and O. falicia, all species presenting a bifurcation in the dorsolateral process of segment IX. Oecetis ruschii sp. nov. can be differentiated by the dorsolateral process of segment IX having a long ventrolateral bifurcation and a short dorsal subapical one, in O. furcata there is a single ventral bifurcation, and in O. falicia it is dorsal and short. The inferior appendage distal lobe in the new species is wider in lateral view than in O. furcata and O. falicia. The apex of this lobe in ventral view is triangular, wide, and flat in the new species while it is regular and digitate in the other species. Additionally, the new species has an asymmetrical projection on the phallic apparatus that is also present in O. falicia but absent in O. furcata.
Adult male: Forewing length 6.2–7.1 mm (n = 5). — Head. Color yellowish brown (in alcohol). Antennae approximately 3x forewing length; scape stout, elongate; pedicel enlarged in width, narrower than scape, shorter than first flagellomeres; first flagellomere narrow, with same length as scape, other flagellomeres shorter than first. Maxillary palps yellowish brown, 5-segmented, segments subequal in length and width, densely covered with setae. Labial palps yellow, apparently 4-segmented, first segment very small. — Thorax. Pterothorax yellowish brown; forewings yellowish brown; dark bands over cord present; dark spots absent; forks I and V rooted (Fig.
The specific epithet is the name of the biological reserve where the specimens were collected, which was named in honor of the naturalist and environmental activist Augusto Ruschi.
In this section we present new distributional records for Oecetis species in Brazil, new records for the country’s states are shown in bold.
BRAZIL • 1♀, 5♂♂; Espírito Santo, Linhares, Lagoa Juparanã Mirim (Lagoa Nova); 19°19′49.0″S, 40°10′12.3″W; 12–13 Sep. 2022; Pennsylvania Light Trap; ADL Viana, P Bonfá Nt, AD Ataíde leg.;
Brazil (Acre, Amazonas, Pará, Bahia, Piauí, Mato Grosso, Rio de Janeiro, São Paulo, Espírito Santo, Minas Gerais), Guiana, Suriname.
BRAZIL • 1♀, 1♂; Minas Gerais, Conceição do Mato Dentro, Cânion do Peixe Tolo; 19°00′15.0″S, 43°36′39.0″W; 01 Dec. 2020; light over white cloth; FF Salles leg.;
Brazil (Minas Gerais, Tocantins)
BRAZIL, • 31♀♀, 1♂; Espírito Santo, Linhares, Lagoa Juparanã Mirin (Lagoa Nova); 19°19′49.0″S, 40°10′12.3″W; 12–13 Sep. 2022; Pennsylvania Light Trap; ADL Viana, P Bonfá Nt, AD Ataíde leg.
Bahamas, Brazil (Amazonas, Bahia, Paraíba, Pernambuco, Piauí, Mato Grosso, Minas Gerais, Rio de Janeiro, São Paulo, Espírito Santo, Paraná, Santa Catarina), Canada, Colombia, Costa Rica, Cuba, El Salvador,United States, Guatemala, Honduras, Jamaica, Mexico, Nicaragua, Panama, Peru, Puerto Rico, Venezuela.
BRAZIL• 1♂; Espírito Santo, Linhares, São Rafael, cachoeira de Angeli; 19°20′55.6″S, 40°25′17.9″W; 25 Sep. 2020; light over white cloth; FF Salles, P Bonfá Nt, TS Raymundo, DS Ferreira leg.;
Argentina, Brazil (Amazonas, Bahia, Pernambuco, Mato Grosso do Sul, Minas Gerais, São Paulo, Espírito Santo), Paraguay, Peru.
Better supported clades within the falicia group are the Nearctic and Central America species (O. arizonica + O. prolongata), the large clade including most South American species with inferior appendage presenting ventral lobe but without dorsal lobe (clades B, C, D, and E), and the clade E formed by species with forked dorsolateral process of segment IX. Most other clades showed low statistical support, suggesting that they can change significantly as more specimens and characters are included in the analysis. The species of the included testacea group (O. akimi, O. testacea, and O. iara) have long autapomorphic branches that might indicate phylogenetic noise. As suggested by
The distribution of the species in clade A suggests that this clade is widespread from southern USA, Central America, and northern South America (Fig.
The historical connections between the Atlantic and Amazon Forests with their expansion over the dry vegetation (Cerrado ecoregion) has been advocated as a general hypothesis to the explanation of disjunct distributions of lineages inhabiting the two ecoregions, with an older connection occurring through a southern route during the Miocene, and a more recent connection in a route through the Northeast Region during the Pliocene and Pleistocene and associated with the Quaternary climate changes (
The asymmetric spiny process on the phallotheca is indicated to appear a single time within the falicia group and lost in O. furcata. An asymmetrical process evolved several times in different species groups as it is present also in the avara and punctata groups. The asymmetry in the phallic apparatus is also a synapomorphy of the inconspicua group, although the asymmetry in this group is not associated with the presence of a spiny process, but on the overall shape of the phallic apparatus.
In this study, we included additional available molecular evidence to the morphologic phylogenetic estimations of the Neotropical Oecetis and a larger sample of Neotropical representatives of the falicia group, including two new species from Atlantic Forest. The main conclusions are:
(1) Most species groups were recovered as monophyletic, however the punctata, pratti and avara groups had low to extremely low supports. In
(2) The two new species from the falicia group herein described have an asymmetric spiny projection on the phalloteca, which was indicated to have evolved a single time in this species group and lost in the O. furcata. Asymmetrical phallic projections also evolved independently in the avara and punctata group.
(3) Most of the falicia group species that share this character occur primarily near the Atlantic coast of South America on the Atlantic Forest and Cerrado ecoregions. A single species is known from the Amazon Forest, and a very disjunct species, O. falicia, is known from Central America. The large gap between O. falicia and related taxa from the Atlantic Forest suggest the existence of undescribed species in this lineage or a wider distribution range of the known species.
(4) The function of the asymmetric spine on male genitalia remains uncertain, and it is not associated with any conspicuous asymmetry in the female internal genitalia. This lack of asymmetry in females supports
The data underlying this article, including the resulting tree (.tre), the morphological (.ss) and the COI matrix (.fas), as well as the combined matrix used in the Bayesian analysis (.nex) are available at the Open Science Framework (OSF) repository and can be accessed at https://osf.io/cp56j/?view_only = b431ca26c7d94ffa9a0ed08376c1cccd
This study was financially supported by the Conselho Nacional de Desenvolvimento Científico (CNPq, grant process 309666/2019–8) and Fundação de Amparo à Pesquisa e Inovação do Espírito Santo (FAPES, grant process 60016604/12; 61938408/13) to FFS; Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – Brasil (CAPES) – Finance Code 001 (006/2022 PROPG-PDEE) to AV; and (CAPES) – Finance Code 001 (process 88887.899774/2023-00) to PBN.
We are grateful to the members of the Laboratory of Systematics and Ecology of Insects (LabSEI, UFES) and the Museu de Entomologia UFV who contributed to collections and field trips. We thank the reviewers and editor for reading and commenting on our manuscript.
Files S1, S2
Data type: .zip
Explanation notes: File S1. Material type and material analyzed standardized in Darwin Core (DWC). — File S2. Additional information from distributional records shown on the map (Fig.
File S3
Data type: .pdf
Explanation notes: Material Morphological tree. Single most parsimonious tree generated at TNT with the adjusted K = 15. Branch support values are displayed near the branches: above Relative Bremer support (from branch-swapping of suboptimal trees up to 10 steps longer and relative fit of 0.9%), below in bold Symmetric Resample (100 replicates, with a change probability of 0.33).