103 urn:lsid:arphahub.com:pub:77d0745d-c3a1-5248-81de-8cdc02bed84a urn:lsid:zoobank.org:pub:F56F6CF9-7502-4001-A751-35D5F2EF6CA0 Arthropod Systematics & Phylogeny ASP 1863-7221 1864-8312 Senckenberg Gesellschaft für Naturforschung 10.3897/asp.84.e154913 154913 Research Article Cerambycidae Chrysomeloidea Coleoptera Insecta Polyphaga Identification key Phylogeny Taxonomy A revised morphology-based phylogeny of Coccoderus Buquet, 1840 (Coleoptera: Cerambycidae: Cerambycinae: Torneutini) and biogeographic analyses reveal diversification on Chacoan dominion Ferreira Gabriel S. gdsferreira1@gmail.com https://orcid.org/0000-0002-4228-9100 1 Conceptualization Writing - original draft Writing - review and editing Data curation Formal analysis Investigation Methodology Roza André Silva https://orcid.org/0000-0003-0886-5159 1 Writing - review and editing Data curation Formal analysis Methodology Validation Mermudes José Ricardo M. https://orcid.org/0000-0003-2030-7483 1 Writing - review and editing Data curation Formal analysis Investigation Methodology Visualization Laboratório de Entomologia, Departamento de Zoologia, Universidade Federal do Rio de Janeiro, Av. Carlos Chagas Filho, 373, Sala A1-107, CEP: 21941-599, Rio de Janeiro, RJ, Brazil Universidade Federal do Rio de Janeiro Rio de Janeiro Brazil https://ror.org/03490as77

Corresponding author: Gabriel S. Ferreira (gdsferreira1@gmail.com)

 2026 06 03 2026 84 235 251 F3A55F68-2E70-55D6-8B58-34E3A886FB48 0 13D902CB-E9EB-40DC-9CD6-D934D63C6D24 04 04 2025 19 12 2025 Gabriel S. Ferreira, André Silva Roza, José Ricardo M. Mermudes 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. https://zoobank.org/13D902CB-E9EB-40DC-9CD6-D934D63C6D24

Abstract

Coccoderus Buquet, 1840 is one of the largest genera in Torneutini (Cerambycinae), composed of 14 species. A previous morphology-based phylogeny recovered the genus as a monophyletic group. Since then, four species were described in the genus. Herein, we expanded this previous morphological dataset with the inclusion of seven characters and eight species. We performed maximum parsimony, maximum likelihood and Bayesian inference analyses. We used the Statistical Dispersal-Vicariance Analysis and the Bayesian Binary Method to reconstruct the potential ancestral ranges of Coccoderus. Our phylogenetic analysis recovered Coccoderus as a monophyletic group and shows shifts in species relationships, with two clades presenting new placements compared to the previous analysis. Biogeographic analyses identified the Chacoan dominion as a key region in the diversification of the genus, representing its ancestral area. Also, we described Coccoderus trimaculatussp. nov., from Brazil, increasing the known species of Coccoderus to 15.

 Taxonomy Evolution longhorn beetles Neotropical 1. Introduction

The tribe Torneutini (Cerambycinae) is exclusively Neotropical, comprising 14 genera and 57 species (Tavakilian and Chevillotte 2025). Coccoderus Buquet, 1840 is one of the largest genera within the Torneutini, comprising 14 species (Tavakilian and Chevillotte 2025). Its distribution is predominantly South American, with only two records in Central America (Costa Rica) (Tavakilian and Chevillotte 2025). Monné (2005) revised the genus Coccoderus, recognizing ten species. Additionally, Monné (2005) conducted phylogenetic analyses based on 31 morphological characters and 12 species (10 Coccoderus species and two outgroup taxa). These analyses resulted in two equally parsimonious trees and recovered Coccoderus as a monophyletic group comprising two distinct clades.

Until Monné (2005), the genus Coccoderus was restricted to South America. In her work, she did not perform formal biogeographical analyses, but discussed the biogeography of the two clades and their respective species in the context of the Neotropical subregions proposed by Morrone (2001; 2004). According to her discussion: (1) Coccoderus species are absent west of the Andes Mountain range and from the Caatinga (northeastern Brazil); (2) species have been recorded in all subregions proposed by Morrone (2001; 2004): Amazonian, Caribbean, Chacoan, and Paraná; and (3) most species of Coccoderus occur in the Chacoan subregion.

Following the taxonomic revision and phylogenetic analysis by Monné (2005), four new species of Coccoderus were subsequently described (Martins & Esteban-Durán 2012; Joly 2017; Ferreira et al. 2023). In addition, several species have been recorded from new localities since Monné (2005), including the expansion of the genus into Central America and the Caatinga (Ferreira et al. 2023; Tavakilian & Chevillotte 2025).

Phylogenetic analyses of the genus Coccoderus using different reconstruction methods (Maximum Parsimony, Maximum Likelihood, and Bayesian Inference) are presented herein. These analyses encompass all known species and incorporate novel morphological characters, the inclusion of new outgroups, and topology comparison tests. Additionally, the first biogeographic analyses for the genus are provided, including ancestral area reconstruction and the formulation of associated biogeographic hypotheses. A new species of Coccoderus is described.

 2. Material and methods 2.1. Taxonomy

The studied specimens were identified by comparison with original descriptions and redescriptions, taxonomic keys (Monné 2005; Jolly 2017; Ferreira et al. 2023), and photographs of type specimens available on Bezark (2025). Photographs were taken using a Leica M205 stereomicroscope, equipped with a DFC450 digital camera, and image stacks were processed using Leica Application Suite v4.3 software (Leica Microsystems). The photographs were edited, and the figure plates were prepared using Adobe Photoshop (Adobe Systems). The terminology follows Lawrence et al. (2010).

The acronyms used in the text are as follows: DZRJ – Coleção Entomológica Professor José Alfredo Pinheiro Dutra, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; MNRJ – Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; USNM – United States National Museum of Natural History, Smithsonian Institution, Washington, DC, United States of America.

 2.2. Phylogenetic analysis

To conduct a phylogenetic analysis aimed at elucidating relationships within the genus Coccoderus, the morphological dataset published by Monné (2005) was used. The ingroup comprised all species currently described in the genus, including the new species described herein, totaling 15 species (Table 1).

List of species of Coccoderus included in the phylogenetic and biogeographical analysis.

Species of Coccoderus
Coccoderus amazonicus Bates, 1870
Coccoderus biguttatus Martins, 1985
Coccoderus bisignatus Buquet, 1840
Coccoderus costae Ferreira, Ferreira & Bravo, 2023
Coccoderus guianensis Tavakilian & Monné, 2002
Coccoderus longespinicornis Fuchs, 1964
Coccoderus novempunctatus (Germar, 1823)
Coccoderus pittieri Joly, 2017
Coccoderus sexguttatus Waterhouse, 1880
Coccoderus sexmaculatus Buquet, 1840
Coccoderus sicki Lane, 1949
Coccoderus speciosus Gounelle, 1909
Coccoderus spinulicornis Joly, 2017
Coccoderus timbaraba Martins & Esteban-Durán, 2012
Coccoderus trimaculatussp. nov.

The outgroup included the two species of Torneutini used by Monné (2005) [Psygmatocerus wagleri Perty, 1828 and Spathopygus eburioides (Blanchard in Orbigny, 1879)], the latter recovered as the sister group of Coccoderus in the previous analyses. In addition, Achryson surinamum (Linnaeus, 1767) [Achrysonini], Chlorida festiva (Linnaeus, 1758) [Bothriospilini], and Retrachydes thoracicus thoracicus (Olivier, 1790) [Trachyderini] were included. Fragoso et al. (1987) proposed two supertribes, Cerambycoinia and Trachyderoinia, based on the structure of the female terminalia. Trachyderoinia included Basipterini, Bothriospilini, Pyrestini (currently Pseudolepturini), Torneutini, and Trachyderini (Fragoso et al. 1987). Given that A. surinamum belongs to Cerambycoinia, this species was used as the outgroup to root the tree topology.

The characters adapted from Monné (2005), as well as the new characters, follow the character statements by Sereno (2007). The matrix was constructed using MESQUITE v3.81 (Maddison and Maddison 2023). The current debate regarding the relative performance of parsimony vs. probabilistic phylogenetic methods when applied to morphological data remains unresolved. Different authors have advocated for the use of parsimony (Goloboff et al. 2008a; Goloboff et al. 2017), probabilistic methods (Wright and Hillis 2014; Puttick et al. 2017; O’Reilly et al. 2018; Barbosa et al. 2024), or even suggested that both approaches yield largely congruent results (Smith et al. 2019). Consequently, all three analyses (Maximum Parsimony, Maximum Likelihood and Bayesian Inference) were performed on our dataset, and their results were briefly compared.

A phylogenetic analysis using Maximum Parsimony (MP) was conducted in TNT v1.6 (Goloboff & Morales 2023), with all characters treated as equally weighted. The analysis was performed under a Traditional Search, using 10,000 replicates and saving 30 trees per replication, with Tree Bisection and Reconnection (TBR). Branch support was assessed using the absolute Bremer (Bremer 1994) and nonparametric bootstrap (Efron 1979; Felsenstein 1985) with 2,000 replicates.

A Maximum Likelihood (ML) analysis was performed in IQTREE2 (Minh et al. 2020), and a Bayesian inference (BI) analysis was conducted in MRBAYES v3.2.7a (Huelsenbeck and Ronquist 2001). Model selection carried out with ModelFinder (Kalyaanamoorthy et al. 2017) in IQTREE2 selected the MKV model (Lewis 2001), with equal state frequencies and correction for ascertainment bias (i.e., MK+FQ+ASC) (see File S1). The BI analysis was run for 50,000,000 generations, saving trees every 1,000 generations and discarding the first 25% as burn-in. Convergence was assessed using TRACER v1.6 (Rambaut et al. 2014).

Node support values were relatively low across all our analyses, which is expected with small matrices (Soltis & Soltis 2003) due to the limited number of characters and a low taxon-to-character ratio (Bremer et al. 1999; Alfaro et al. 2003; Erixon et al. 2003; Wiesemüller & Rothe 2006). We followed the thresholds proposed by Zander (2004) to interpret support values in morphological datasets: high support corresponds to Bootstrap values above 88%, Bremer support above 3, and Posterior Probability (PP) above 91%. For UFBoot values, we followed the commonly accepted threshold of ~95% for strong support (Hoang et al. 2018).

We refrain from using fixed thresholds as hard cutoffs, as this could be overly restrictive given the limited empirical basis for defining support levels in morphological analyses. To account for this, we also defined a “medium” support range (except for Bremer support), which is the range of 70–95% for UFBoot, and 70–91% for Posterior Probability. The lower threshold of 70% was adopted based on the classical study of Hillis & Bull (1993) which established this approximated value under specific conditions that has since been informally used as a “rule of thumb” value in phylogenetic support interpretation.

To compare the topologies of the phylogenetic trees, the Robinson–Foulds (RF) distance test (Robinson & Foulds 1981) was performed in IQTREE2 (Minh et al. 2020), which quantifies the difference between two trees based on the number of distinct bipartitions. The resulting distances were then normalized to a range from 0 to 1 using the statistical program R version 4.3.3. Trees with mapped synapomorphies were visualized using the WinClada-ASADO version (Nixon 2004), with unambiguous optimization.

 2.3. Biogeographic analysis

The distribution of Coccoderus species was classified into six biogeographic areas, following the dominions and zones defined by Morrone (2014; 2022): A (Pacific Dominion), B (Boreal Brazilian Dominion), C (South Brazilian Dominion), D (Southeastern Amazonian Dominion), E (Chacoan Dominion and Paraná Dominion), and F (South American Transition Zone). The Paraná Dominion was grouped with the Chacoan Dominion to decrease the number of areas and simplify the analysis, as species occurring in the former are always present in the latter, but the reciprocal is not true. The information about their distribution was based on recent works (Monné 2005; Jolly 2017; Ferreira et al. 2023; Tavakilian & Chevillotte 2025). Geographical coordinates, when not available on specimen labels, were obtained using Google Maps (www.google.com/maps). A distribution map was generated with QUANTUM-GIS 3.34.12 (QGIS Development Team 2024) using the shapefile of dominions provided by Morrone et al. (2022).

We used Statistical Dispersal-Vicariance Analysis (S-DIVA) (Yu et al. 2010) and Bayesian Binary Method (BBM) (Ronquist and Huelsenbeck 2003), both implemented in RASP 4.0 (Yu et al. 2015), to reconstruct the potential ancestral ranges of Coccoderus on the phylogenetic trees. We chose these two methods because both allow reconstructions using trees lacking branch lengths, enabling us to select any of the topologies from the phylogenetic analyses for ancestral area reconstruction. To account for phylogenetic uncertainty, we analyzed 3,192 trees sampled from the MCMC output and ran both methods. S-DIVA was performed allowing for extinction events. BBM was run under a fixed state frequency model (Jukes–Cantor) (Jukes & Cantor 1969) with equal among-site rate variation, over two million generations and ten independent chains.

The maximum number of areas per node was set to six, matching the number of dominions in which the species occur, thus allowing for all possible combinations of ancestral areas (Kodandaramaiah 2010). The closest outgroup taxa (or clade, when applicable) was retained in the biogeographical analysis to minimize bias towards inferring a widespread ancestor at the root (Ronquist 1997; Kodandaramaiah op. cit.).

 3. Results 3.1. Taxonomy

CERAMBYCIDAE Latreille, 1802

CERAMBYCINAE Latreille, 1802

TORNEUTINI Thomson, 1861

Coccoderus Buquet, 1840

Animalia Coleoptera Cerambycidae 83B3CFE6-3DC5-57DC-AF8D-711ACE47761A Coccoderus trimaculatus https://zoobank.org/1E8336E3-DF09-4900-B858-7C13B324EF9E sp. nov. Figure 1A–E Description.

Holotype female. Coloration. Integument mostly orangish brown; apex of antennomeres reddish brown, inner area of mandible, pronotal tubercles, and lateral tubercles of prothorax black; elytra transparent with eburneous callosities yellowish white; tarsal claws dark brown. — Head. Frons, clypeus, labrum, mandibles, labial and maxillary palpi, and antennae with sparse, long, and erect yellowish setae. Antennal tubercles and vertex with thick, shallow, dense punctation. Antennal tubercles globose, well separated; with shallow sulcus between antennal tubercles. Upper eye lobes well separated, distance between them about 2.8 times the width of one upper lobe; upper eye lobe with nine rows of ommatidia dorsally. Mandible with one tooth on the inner margin. Genae acuminated. Antennae with 12 antennomeres, not reaching elytral apex; external apex of antennomeres III–VI with short spiniform projection; antennomere III about twice the length of IV; antennomeres IV–X with subequal length; XI–XII longer than the previous. — Thorax. Prothorax wider than long; with thick, shallow, and dense punctuation; each side of the prothorax with two rounded tubercles. Prothorax subglabrous, with rows of dense and erect setae on anterior and posterior margins. Pronotum with two elevated tubercles before the middle, rounded apically. Prosternum and mesoventrite with short and erect yellowish setae. Prosternal process rounded apically. Mesosternal process truncated. Metaventrite covered with dense, long, and decumbent yellowish pubescence, glabrous metathoracic discrimen. — Elytra. Each elytron with three elliptical eburneous callosities, with different lengths, one anteriorly, one medial, and another one on posterior third; distance between anterior and medial callosities and between medial and posterior callosities subequal to length of medial callosity. Elytral apex with external and sutural spine. — Legs. With sparse, long, and erect yellowish setae. Femora subfusiform; apex of meso- and metafemora with spine on inner margin. Tibia cylindrical, as long as femora. — Abdomen. Ventrites covered with dense, long, and decumbent yellowish pubescence; ventrite 5 with apical margin emarginate.

10.3897/asp.84.e154913.figure1 C176440E-7CB8-5323-8452-F21FF26F3335

A–ECoccoderus trimaculatussp. nov., holotype female. A dorsal view; B ventral view; C lateral view; D frontal view; E prothorax.

https://binary.pensoft.net/fig/1555952 Variation.

In the paratype, the integument is mostly yellowish brown. The distance between upper eyes lobes is about 2 times the width of one upper lobe. Distance between anterior and medial callosities about 1.2 times the length of medial callosity; distance between medial and posterior callosities about 1.1 times the length of medial callosity.

 Measurements.

(in mm) Holotype female, total length: 35.0; prothorax length: 5.3; prothorax width: 6.8; anterior width of prothorax: 5.5; posterior width of prothorax: 6.4; elytral length: 25.6; humeral width: 8.4. Paratype female, total length: 33.6; prothorax length: 5.7; prothorax width: 7.3; anterior width of prothorax: 5.5; posterior width of prothorax: 6.8; elytral length: 25.2; humeral width: 8.0.

 Etymology.

Latin, “tri” (three) and “maculatus” (spots); allusive to the elytra with three pairs of eburneous callosities.

 Type material.

Holotype female, BRAZIL, São Paulo, [Ubatuba] Picinguaba, 03–07.xi.2010, Mattos, I. & Mermudes, J.R.M. col. (DZRJ) Paratype female, BRAZIL, Minas Gerais, Jaboticatubas, Parque Nacional da Serra do Cipó, Alojamento dos Brigadistas, 19°21’0.1”S 43°37’07.4”W, 809 m, 17.xi.2018, pano branco, Alves, A.A. col. (MNRJ)

 Remarks.

Coccoderus trimaculatussp. nov. is morphologically similar to Coccoderus costae Ferreira, Ferreira & Bravo, 2023 by the presence of large eburneous callosities on elytra, antennomere III with spiniform projection on outer margin, and more than twice as long as IV and femora subfusiform. However, Coccoderus trimaculatussp. nov. has the upper eye lobe with nine rows of ommatidia dorsally; antennomere III about 2 times the length of IV; eburneous callosities yellowish white; distance between medial and posterior callosities subequal to length of medial callosity. It differs from C. costae by the upper eye lobe with eight rows of ommatidia dorsally; antennomere III about 3 times the length of IV; eburneous callosities on elytra yellowish; distance between medial and posterior callosities about one-third the length of the medial callosity. Also differs from females of C. sexmaculatus by the upper eye lobe with nine rows of ommatidia dorsally (rather than eight rows of ommatidia) and elytra transparent (rather than opaque)

 Distribution.

Coccoderus trimaculatussp. nov. is recorded so far only in the type locality, in São Paulo and Minas Gerais, both in Brazil, in the Brazilian biome of the Atlantic Forest, C. costae is only recorded in the Brazilian state of Bahia (Brazil), in the Brazilian biome of the Caatinga, and C. sexmaculatus occurs in the Brazilian states of Goiás, Distrito Federal, Bahia, Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo, in the Brazilian biomes of the Atlantic Forest and Cerrado.

 3.2. New records Animalia Coleoptera Cerambycidae E13B05C9-750C-57F7-BAC4-548A83CB3A52 Coccoderus amazonicus Bates, 1870 Distribution.

Costa Rica, Ecuador, Colombia, Venezuela, Peru, Suriname, Brazil (Amazonas, Pará, Minas Gerais), Bolivia (Santa Cruz).

 Material examined.

1 female, SURINAME [New country record], Brokopondo District, Brownsberg Natuurpark, Mazaroni Plateau, 400–500m, 17 August 1982, Collins, Barly, Oberman, Pollock, Putnam, Steiner col (USNM). 1 female, SURINAME, Brokopondo District, Brownsberg Natuurpark, Mazaroni Plateau, 400–500m, 21 August 1982, Collins, Barly, Oberman, Pollock, Putnam, Steiner col (USNM).

Key to the species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Coccoderus">Coccoderus</tp:taxon-name-part></tp:taxon-name></italic> Buquet, 1840

(modified from Ferreira et al (2023))

1 Elytra without eburneous callosities. BRAZIL (Maranhão, Rio Grande do Norte, Bahia, Mato Grosso, Goiás, Distrito Federal, Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, and Rio Grande do Sul), BOLIVIA (Santa Cruz, Tarija), PARAGUAY, ARGENTINA (Misiones, Salta, Santiago del Estero, Chaco, Entre Ríos, and Buenos Aires), and URUGUAY Coccoderus novempunctatus (German, 1823)
1’ Elytra with eburneous callosities 2
2 (1). — Elytra with only one pair of eburneous callosities, located between humeri and scutellum 3
2’ Elytra with three pairs of eburneous callosities, located along the elytra 4
3 (2). Antennomere III with outer apical spine; prosternal process without tubercle. BRAZIL (Mato Grosso) and BOLIVIA (Santa Cruz) Coccoderus biguttatus Martins, 1985
3’ Antennomere III without outer apical spine; prosternal process with tubercle. VENEZUELA, SURINAME, FRENCH GUIANA, BRAZIL (Amapá, Pará, and Maranhão) Coccoderus bisignatus Buquet, 1840
4 (2). — Meso- and metafemur without long spine on inner apex; elytral apex with only sutural spine. BRAZIL (Maranhão, Bahia, Mato Grosso, Goiás, Distrito Federal, Minas Gerais, Mato Grosso do Sul, and São Paulo) Coccoderus speciosus Gounelle, 1909
4’ Meso- and metafemur with long spine on inner apex; elytral apex with lateral and sutural spines 5
5 (4). — Elytra opaque or elytra opaque and slightly translucid 6
5’ Elytra translucid 7
6 (5). — Head with tick and dense punctation; elytra opaque. BRAZIL (Bahia, Goiás, Distrito Federal, Minas Gerais, Espírito Santo, Rio de Janeiro, and São Paulo). . . . Coccoderus sexmaculatus Buquet, 1840
6’ Head with thin and moderately dense punctation; elytra opaque and slightly translucid. VENEZUELA Coccoderus spinulicornis Joly, 2017
7 (5’). — Elytra with eburneous callosities shorter than half of antennomere III 8
7’ Elytra with eburneous callosities longer than half of the antennomere III 10
8 (7). — Pronotum with thick, deep and dense punctation. VENEZUELA, FRENCH GUIANA, PERU, BRAZIL (Amapá, Amazonas, Pará, Maranhão, Piauí, Rondônia, Mato Grosso, Goiás, Distrito Federal, Minas Gerais, and Mato Grosso do Sul) and PARAGUAY Coccoderus longespinicornis Fuchs, 1964
8’ Pronotum with irregular or thin, shallow and dense punctation 9
9 (8’). — Mandible with tooth on the inner margin (male); antennomere III with long and dense setae. COSTA RICA, ECUADOR, COLOMBIA, VENEZUELA, SURINAME, BRAZIL (Amazonas, Pará, and Minas Gerais), PERU and BOLIVIA (Santa Cruz) Coccoderus amazonicus Bates, 1870
9’ Mandible falciform, without tooth on the inner margin (male); antennomere III with short and sparse setae. FRENCH GUIANA and BRAZIL (Amapá) Coccoderus guianensis Tavakilian & Monné, 2002
10 (7’). — Antennomere III with rounded projection on external apex (except, females of Coccoderus sexguttatus Waterhouse, 1880 with spiniform projection) 11
19’ Antennomere III with spiniform projection on external apex 12
11 (10). — Antennal tubercles separated by deep sulcus; apical spine of meso- and metafemora almost cylindrical on apical half Coccoderus pittieri Jolly, 2017
11’ Antennal tubercles not separated by deep sulcus; apical spine of meso- and metafemora more triangular. COLOMBIA, VENEZUELA, ECUADOR and PERU Coccoderus sexguttatus Waterhouse, 1880
12 (10’). — Female. Antennomere III slightly longer than IV; antennomeres IV–XII decrescent in length. BRAZIL (Amapá, Pará, Rondônia, Mato Grosso, Goiás, and Distrito Federal) Coccoderus sicki Lane, 1949
12’ Female. Antennomere III more than twice longer than IV; antennomeres IV–XI subequal in length 13
13 (12’). — Female. Eburneous callosities on elytra subequal in length; prosternal process truncated apically. COSTA RICA Coccoderus timbaraba Martins & Duran, 2012
13’ Female. Eburneous callosities on elytra with different lengths; prosternal process rounded apically 14
14 (13’) Female. Upper eye lobe with eight rows of ommatidia dorsally; antennomere III about 3× the length of IV; eburneous callosities on elytra yellowish; distance between medial and posterior callosities about one-third length of medial callosity. BRAZIL (Bahia) Coccoderus costae Ferreira, Ferreira & Bravo, 2023
14’ Female. Upper eye lobe with nine rows of ommatidia dorsally; antennomere III about 2× the length of IV; eburneous callosities whitish (Fig. 1A); distance between medial and posterior callosities subequal to length of medial callosity (Fig. 1A). BRAZIL (Minas Gerais and São Paulo) Coccoderus trimaculatus sp. nov.
3.3. Morphological characters

Thirty-eight morphological characters were used: nine from the head, 11 from the prothorax, seven from the elytra, six from the legs, one from the abdomen, three from the male terminalia, and one from the female terminalia (see File S2). Of these, seven are new characters, 30 were modified from Monné (2005), and one was modified from Monné & Napp (2005). For each character, we provide the number of steps (L), the consistency index (CI), and the retention index (RI).

Head

1. Males, lower eye lobes, narrow extension to the ventral region: (0) absent; (1) present (modified and translated from Monné 2005, fig. 58). L = 1; ci = 1; ri = 1.

2. Female, mandible, region between the base and the median tooth: (0) not excavated; (1) excavated (modified and translated from Monné 2005, figs 11, 12). L = 1; ci = 1; ri = 1.

3. Maxillary palpi, shape of the palpomeres: (0) conical; (1) moderately conical; (2) cylindrical (modified and translated from Monné 2005). L = 3; ci = 0.66; ri = 0.50.

4. Maxillary palpi, length of the palpomeres II–III: subequal (0); II longer than III (1); II shorter than III (2) (modified and translated from Monné 2005). L = 4/5; ci = 0.50/0.40; ri = 0.75/0.62.

5. Antennae, number of antennomeres: (0) 11 antennomeres; (1) 12 antennomeres. L = 1; ci = 1; ri = 1.

6. Antennomere III, brush of setae: (0) absent; (1) present (translated from Monné 2005). L = 3/2; ci = 0.33/0.50; ri = 0.50/0.75.

7. Male, antennomere III, length relative to IV: (0) III longer than IV; (1) III shorter than IV (translated from Monné 2005). L = 2; ci = 0.50; ri = 0.50.

8. Antennomere III, spiniform projection: (0) absent; (1) present (modified and translated from Monné 2005). L = 3; ci = 0.33; ri = 0.71.

9. Antennomere III, spiniform projection, length: (0) shorter than the width of the antennomere; (1) longer than the width of the antennomere (modified and translated from Monné 2005). L = 2; ci = 0.50; ri = 0.50.

Thorax

10. Sides of prothorax, black spots: (0) absent; (1) present (translated from Monné 2005). L = 1; ci = 1; ri = 1.

11. Sides of prothorax, antemedian tubercles: (0) absent; (1) present (1) (modified and translated from Monné 2005). L = 4; ci = 0.25; ri = 0.

12. Pronotum, microsculpted punctuation: (0) absent; (1) present (translated from Monné 2005). L = 1; ci = 1; ri = 1.

13. Pronotum, anterior region, rounded spots: (0) absent; (1) present. L = 1; ci = 1; ri = 1.

14. Pronotum, anterior region, length of spots: (0) shorter or equal to the width of the pedicel; (1) longer than width of the pedicel. L = 2; ci = 0.50; ri = 0.50.

15. Pronotum, anterior margin: (0) truncated; (1) rounded; (2) rounded and slightly sinuous (modified and translated from Monné 2005). L = 3; ci = 0.66; ri = 0.87.

16. Pronotum, lateroposterior tubercles: (0) absent; (1) present (translated from Monné 2005). L = 2; ci = 0.50; ri = 0.80.

17. Prosternum, shape: (0) flat; (1) depressed; (2) swollen (modified and translated from Monné 2005). L = 3; ci = 0.66; ri = 0.50.

18. Prosternum, coxal cavity: (0) non-angular; (1) slightly angular; (2) strongly angular (modified from Monné & Napp 2005, figs 65–70). L = 2; ci = 1; ri = 1.

19. Prosternal process, apical tubercle: (0) absent; (1) present (modified and translated from Monné 2005). L = 2; ci = 0.50; ri = 0.83.

20. Mesosternal process, shape: flat (0); convex (1) (modified and translated from Monné 2005). L = 2; ci = 0.50; ri = 0.75.

Elytra

21. Elytra, setae: (0) absent; (1) present. L = 2; ci = 0.50; ri = 0.66.

22. Elytra: (0) transparent; (1) opaque. L = 3; ci = 0.33; ri = 0.66.

23. Elytra, region between the humeri and the scutellum, eburneous callosities: (0) absent; (1) present (modified and translated from Monné 2005). L = 2; ci = 0.50; ri = 0.75.

24. Elytra, medial region, eburneous callosities: (0) absent; (1) present. L = 4; ci = 0.25; ri = 0.50.

25. Elytra, posterior region, eburneous callosities: (0) absent; (1) present. L = 4; ci = 0.25; ri = 0.50.

26. Elytra, latero-basal depression: (0) absent; (1) present (translated from Monné 2005). L = 1; ci = 1; ri = 1.

27. Elytral apex, lateral spine: (0) absent; (1) present (translated from Monné 2005). L = 3; ci = 0.33; ri = 0.33.

Legs

28. Internal apex of mesofemora, spine: (0) absent; (1) present (modified and translated from Monné 2005). L = 4; ci = 0.25; ri = 0.25.

29. Internal apex of mesofemora, acuminate projection: (0) absent; (1) present (modified and translated from Monné 2005). L = 3; ci = 0.33; ri = 0.33.

30. Metafemora, shape: linear (0); subfusiform (1); fusiform (2) (modified and translated from Monné 2005). L = 3; ci = 0.66; ri = 0.83.

31. Metafemora, ventral region, brush of setae: (0) absent; (1) present (modified and translated from Monné 2005). L = 2; ci = 0.50; ri = 0.

32. Internal apex of metafemora, spine: (0) absent; (1) present (modified and translated from Monné 2005). L = 3; ci = 0.33; ri = 0.

33. Internal apex of metafemora, acuminate projection: (0) absent; (1) present (modified and translated from Monné 2005). L = 2; ci = 0.50; ri = 0.75.

Abdomen

34. Ventrite 5, apical margin: emarginated (0); truncated (1); rounded (2) (modified and translated from Monné 2005). L = 4; ci = 0.50; ri = 0.60.

Male terminalia

35. Sternite VIII, apical margin, lobes: (0) absent; (1) present (modified and translated from Monné, 2005, figs 31, 33). L = 3; ci = 0.33; ri = 0.33.

36. Sternite IX, apophysis, length in relationship to the length of rest of sternite: (0) shorter; (1) longer (modified and translated from Monné 2005, figs 36–38). L = 2; ci = 0.50; ri = 0.75.

37. Sternite IX, apex: (0) straight; (1) curved (modified and translated from Monné, 2005, figs 37, 38). L = 2; ci = 0.50; ri = 0.50.

Female terminalia

38. Sternite VIII, apophysis, apex: (0) narrow; (1) apex about three times wider than the median region; (2) apex about four times wider than the median region (translated from Monné 2005, figs 62–64, 69, 72, 75, 78). L = 3; ci = 0.66; ri = 0.75.

 3.4. Phylogenetic analyses

All analyses (MP, ML, and BI), conducted with 20 taxa and 38 characters recovered Coccoderus as a monophyletic group (Figs 2, 3; Files S4, S5). Some clades were consistently recovered across all analyses, such as: (C. amazonicus + C. guianensis), (C. biguttatus + C. speciosus), and (C. longespinicornis + C. spinulicornis) (Figs 2, 3; Files S4, S5).

10.3897/asp.84.e154913.figure2 BA432F5B-9185-5F4F-B293-896BDD3CF222

A, B Phylogenetic hypothesis of Coccoderus on the equal weights Maximum Parsimony analysis selected by the Robinson-Foulds distance test. A First tree; B Clade B of the second tree. The numbers above the clades represent the relative Bremer support and below the clades the nonparametric bootstrap support (values > 70).

https://binary.pensoft.net/fig/1555953 10.3897/asp.84.e154913.figure3 2D65D0CA-8822-5CD2-A03D-E00C57D7BB9E

A, B Phylogenetic relationship of Coccoderus found on the two most parsimonious trees recovered with equal weights parsimony analysis selected by the Robinson-Foulds distance test, First tree, with L= 93 IC= 0.495 IR= 0.659; Second tree, with L=93 CI=0.495 RI=0.659. A First tree; B Second tree. Non-homoplastic synapomorphies are marked with black spots, while homoplastic ones are marked with white spots.

https://binary.pensoft.net/fig/1555954

The Robinson –Foulds distance analysis indicated that trees 2 and 4 from the MP analysis under equal weight were the most similar, showing the smallest distance (Table 2; File S3). The MP analysis with equal weight resulted in four equally parsimonious trees, with 93 steps, the consistency index of 0.495, and the retention index of 0.659. Coccoderus was recovered as monophyletic, supported by Bremer (3) and Nonparametric Bootstrap (60) (Fig. 2). The genus is supported by the following synapomorphies: sides of the prothorax with black spots (10:1, non-homoplastic), anterior region of the pronotum with rounded spots (13:1, non-homoplastic), lateroposterior of the pronotum with tubercles (16:1, non-homoplastic), elytra without setae (21:0, homoplastic), transparent elytra (22:0, non-homoplastic) (Fig. 3).

Normalized Robinson-Foulds distance test. Legend: Bayesian inference (BI), MP= Maximum Parsimony, and Maximum Likelihood (ML).

MP1 MP2 MP3 MP4 BI ML
MP1 0 0.053 0.132 0.053 0.079 1
MP2 0.053 0 0.105 0.026 0.105 1
MP3 0.132 0.105 0 0.105 0.158 1
MP4 0.053 0.026 0.105 0 0.105 1
BI 0.079 0.105 0.158 0.105 0 1
ML 1 1 1 1 1 0

The internal relationships within Coccoderus comprises two clades (Figs 2, 3). The first clade (Clade A) is supported by two synapomorphies: males with apophysis of sternite IX longer than remaining portion of the sternite (36:1, homoplastic) and females with the apex of apophysis of sternite VIII about four times wider than the median region (38:2, non-homoplastic) (Fig. 3), with the support of Bremer = 1 (Fig. 2). The second clade (Clade B) is supported by the rounded anterior margin of pronotum (15:1, homoplastic) (Fig. 3), with the support of Bremer = 1 (Fig. 2). The only difference between the two trees selected by RF distance test concerns the position of C. novempunctatus, within Clade B. In the first tree, C. novempunctatus is recovered as the sister group to the remaining species of this clade, except C. timbaraba (Fig. 3A). In the second tree, C. novempunctatus is recovered as a sister group to the other Coccoderus of this clade, except C. timbaraba and the clade (C. pittieri + C. bisignatus) (Fig. 3B). In both trees, Coccoderus trimaculatussp. nov. is recovered as a sister group of C. costae (Fig. 3).

The Maximum Likelihood analysis recovered Coccoderus as monophyletic, with moderate support of the Ultrafast bootstrap (88%) (File S4). The clade (C. biguttatus + C. speciosus) was recovered as the sister group to the remaining Coccoderus species (File S4). The remaining species were recovered in a polytomy that included C. timbaraba and two clades: (C. sexguttatus + C. pittieri + (C. sicki + (C. longespinicornis + C. spinulicornis))) and (C. costae (C. trimaculatussp. nov. (C. amazonicus + C. guianensis) (C. sexmaculatus + (C. bisignatus + C. novempunctatus)))) (File S3).

Coccoderus was recovered as monophyletic in the Bayesian Inference analysis, with posterior probability support of 0.88 (File S5). The clade (C. biguttatus + C. speciosus) was recovered as a sister group to the remaining Coccoderus species. The clades (C. amazonicus + C. guianensis) and (C. biguttatus + C. speciosus) showed posterior probability above 0.95 (File S5).

 3.5. Biogeographic analyses

The distribution map (Fig. 4) shows that most species are widespread, occurring in two or more dominions, except for species known from one or a few specimens, such as C. costae, C. guianensis, C. pittieri, C. spinulicornis, C. timbaraba, and C. trimaculatus. The BBM and S-DIVA analyses were performed using the topologies obtained by the MP analysis (Figs 5, 6, 7, 8), which resulted from the Robinson–Foulds distance test. Both parsimony topologies were used to account for variation in the internal relationships of Coccoderus.

10.3897/asp.84.e154913.figure4 0CBADA74-BF50-5095-9ECF-1F0490C2FCBC

A, B. Distribution of Coccoderus species: A Distribution of C. biguttatus, C. longespinicornis, C. sexguttatus, C. sicki, C. speciosus, C. spinulicornis, C. pittieri. B Distribution of C. amazonicus, C. bisignatus, C. costae, C. guianensis, C. novempunctatus, C. sexmaculatus, C. timbaraba, C. trimaculatussp. nov.

https://binary.pensoft.net/fig/1555955 10.3897/asp.84.e154913.figure5 BC1B63D5-E518-577A-9A38-8B42F982830F

Ancestral area reconstruction by S-DIVA analysis of the first most parsimonious tree recovered: Letters representing following areas: A Pacific, B Boreal Brazilian, C South Brazilian, D South-eastern Amazonian, E Chacoan and Paraná, and F (South American transition).

https://binary.pensoft.net/fig/1555956 10.3897/asp.84.e154913.figure6 B55D1C0B-3410-5147-B40E-CC2D947E2379

Ancestral area reconstruction by S-DIVA analysis of the second most parsimonious tree recovered: Letters representing following areas: A Pacific, B Boreal Brazilian, C South Brazilian, D South-eastern Amazonian, E Chacoan and Paraná, and F South American transition.

https://binary.pensoft.net/fig/1555957 10.3897/asp.84.e154913.figure7 F8CCCB89-B7C9-5709-8F73-1BCF629DD106

Ancestral area reconstruction by BMM analysis of the first most parsimonious tree recovered: Letters representing following areas: A Pacific, B Boreal Brazilian, C South Brazilian, D South-eastern Amazonian, E Chacoan and Paraná, and F South American transition).

https://binary.pensoft.net/fig/1555958 10.3897/asp.84.e154913.figure8 EAF8C51E-8C1B-5101-AAD4-C358A91DC81B

Ancestral area reconstruction by BMM analysis of the second most parsimonious tree recovered: Letters representing following areas: A Pacific, B Boreal Brazilian, C South Brazilian, D South-eastern Amazonian, E Chacoan and Paraná, and F (South American transition).

https://binary.pensoft.net/fig/1555959

The S-DIVA analysis of the first topology inferred 31 dispersal, eight vicariance, and three extinction events (Fig. 5). The most likely ancestral area of Coccoderus (node 30) was recovered as the composite area ABCDE, with marginal probability of 39%. The second most likely ancestral area was a composed area of all areas (ABCDEF), with 25% probability. The most probable ancestral area of Clade A (node 29) was recovered as area E, with 61% probability. The clade comprising C. biguttatus + C. speciosus (node 25) had area E recovered as ancestral, with 93% probability. The clade formed by (C. longespinicornis + (C. sexguttatus + (C. spinulicornis + C. sicki))) (node 28) had area A recovered as the ancestral area with 31% probability, followed by area E with 19% probability. The following node (27) had area E with 51% probability, and the terminal node of this clade (26) was highly unresolved, with possible ancestral areas are ACDE, ACE, ADE, and AE, all around 17% probability.

On the other hand, Clade B had its ancestral area (Node 23) poorly resolved, with area AE (21% probability) followed by area A (19% probability), along with several composite areas showing lower probabilities. All the other nodes within this clade had poorly resolved ancestral area reconstructions, with multiple different areas competing with similar probability (none adove 30% probability), except for the following: (C. sexmaculatus + ((C. amazonicus + C. guianensis) + (C. costae + C. trimaculatus)) (node 20) with area E (55% probability); ((C. amazonicus + C. guianensis) + (C. costae + C. trimaculatus))) (node 19) with ancestral area BEF (31% probability); (C. amazonicus + C. guianensis) (node 17) with ancestral area B (30% probability); and (C. costae + C. trimaculatus) (node 18) with ancestral area EF (100% probability).

The S-DIVA analysis of the second topology inferred 32 dispersal, eight vicariance, and three extinction events (Fig. 6). The most likely ancestral area of Coccoderus (node 30) was recovered similarly to the first topology, as the composite area ABCDE, with a marginal probability of 36%. The second most likely ancestral area was a composed area of all areas (ABCDEF), with 24% probability. The most probable ancestral area of Clade A (node 29) was recovered as area E, with 43% probability. The clade comprising C. biguttatus + C. speciosus (node 25) had area E as ancestral recovered with 94% probability. The clade formed by (C. longespinicornis + (C. sexguttatus + (C. spinulicornis + C. sicki) (node 28) had area A as ancestral with 29% probability, followed by area E with 20% probability. The subsequent node (27) has area E with 52% probability, and the terminal node of this clade (26) was highly unresolved, with the possibility of an ancestral area of ACDE, ACE, ADE and AE, all around 17% probability.

In contrast, Clade B had its ancestral area (Node 23) poorly resolved, with area AE (21% probability) followed by area A (19% probability), alongside with many other composed areas with smaller probabilities. All other nodes within this clade had their ancestral area poorly reconstructed, with many different areas competing in close probability (none adobe 30% probability), except for the following: (C. novempunctatus + (C. sexmaculatus + ((C. amazonicus + C. guianensis) + (C. costae + C. trimaculatus))) (node 21), with area E (87% probability); (C. sexmaculatus + ((C. amazonicus + C. guianensis) + (C. costae + C. trimaculatus)) (node 20), with area E (41% probability); (C. amazonicus + C. guianensis) (node 17) with ancestral area B (38% probability); and (C. costae + C. trimaculatus) (node 18), with ancestral area EF (100% probability).

The BMM analysis of the first topology resulted in the possibility of 34 dispersal and six vicariance events (Fig. 7). The most likely ancestral area of Coccoderus (node 30) was recovered as area E (Chacoan Dominion), with 51% marginal probability. The most probable ancestral area of Clade A (node 29) was also recovered as area E, with 38% probability, followed closely by area CE (30%). The clade comprising C. biguttatus + C. speciosus had ancestral area E recovered as ancestral, with 67% probability. The clade formed by (C. longespinicornis + (C. sexguttatus + (C. sicki + C. spinulicornis)) (node 28) had area ACE recovered as ancestral area recovered, with 32%. All the other nodes of this clade (Node 26 and 27) also had area ACE as the most probable, although with only 22% probability and closely followed by other composite areas.

Clade B also had its ancestral area inferred as area E, with 33% probability (Node 24). All the other nodes in this clade had area E as the most probable ancestral area (although poorly supported in nodes 22 and 23), with exception of C. amazonicus + C. guianensis (ancestral area B) and C. bisignatus + C. pittieri (several ancestral areas with similar probabilities below 20%).

The second MP topology (Fig. 8) yielded results that differed from those of the first one (Fig. 7) and more similar to those obtained in the S-DIVA analysis of the second topology. The most likely ancestral area of Coccoderus (node 30) was recovered as the composed area ABCDE, with 37% marginal probability. The second most likely ancestral area was a composed area of all areas (ABCDEF), with 24% probability. Clade A showed very similar results in comparison to BBM topology one analysis, regarding ancestral areas recovered and probabilities. The main differences were in node 28 (ancestral area A with 29% probability), node 27 with area E (52% probability), and node 26, which was poorly resolved. Clade B has area AE recovered as more probable (Node 23), with 25% probability, and the following nodes (21 and 22) had area E as the most likely ancestral area (87% and 40% probability, respectively). The nodes 19, 22 and 23 were poorly resolved, and the two last nodes had the following recovered areas: (C. amazonicus + C. guianensis) (node 17) with ancestral area B (37% probability); and (C. costae + C. trimaculatus) (node 18) with ancestral area EF (100% probability). All ancestral area probabilities, as well as recovered possible dispersal and vicariant events are accessible in File S6.

 4. Discussion 4.1. Phylogenetic analyses

All of our analyses recovered the genus Coccoderus as a monophyletic group, but with different topologies. The Maximum parsimony analysis recovered four equally parsimonious trees with high resolution, but the Robinson –Foulds distance analysis showed that trees 2 and 4 were the most similar. The Maximum Likelihood analysis resulted in a single tree, but the internal relationships within the genus remained poorly resolved, with several polytomies. The Bayesian inference analysis showed a low support for some clades within Coccoderus, except for (C. amazonicus + C. guianensis) and (C. biguttatus + C. speciosus), with high support. The low resolution of the ML tree, however, was unexpected and may indicate that relationships within the genus remain far from resolved. Further analyses, potentially including molecular data, are necessary to achieve a more stable topology across different analyses.

This instability is underscored by the substantial differences between the Maximum Parsimony (MP) and Maximum Likelihood (ML) analyses: (1) C. biguttatus and C. speciosus are recovered as the sister group to the remaining species of Clade A in the MP analysis, whereas in the ML analysis, they are recovered as the sister group to all other Coccoderus species; (2) C. timbaraba is the sister to the other species of Clade B in the MP analysis, but is part of a polytomy with species from both Clades A and B in the ML analysis.

Monné (2005) analyzed 12 taxa, including 10 species of Coccoderus, and 31 morphological characters, and performed a Maximum parsimony analysis under equal weighting, which resulted in two equally parsimonious trees. With the inclusion of species described after the revision and phylogenetic analysis of the genus by Monné (2005), some relationships have changed. The topology in Clade A is similar in both parsimony trees, with (C. longespinicornis as the sister group to (C. sexguttatus + (C. spinulicornis + C. sicki)), which differs from Monné (2005), where C. longespinicornis was recovered as the sister group to (C. sexguttatus + C. sicki). In Clade B, C. amazonicus and C. guianensis were recovered as a sister group, differing from Monné (2005), where C. guianensis was recovered either as the sister to C. amazonicus or as the sister to the clade (C. amazonicus + C. sexmaculatus).

 4.2. Biogeographic analyses

The biogeographic analyses were mostly poorly resolved, especially concerning the ancestral area of the genus. Nevertheless, one biogeographic area seems to have played a significant role in the biogeographical history of the genus: The Chacoan Dominion (Area E). The Chacoan Dominion was most likely the ancestral area of the clade A in all analyses, and also the clade B in the BBM topology one. Many of the internal nodes had this area recovered as the most probable ancestral area, so this dominion possibly played a significant role in the diversification of both internal clades. This idea was already pointed out by Monné (2005), who highlighted the amount of species sympatry occurring in this region.

It is interesting to point out that the genus has a very high dispersal ability. All the analyses recovered a large number (above 30) of dispersal events, and several species of both internal clades occur in sympatry in many different dominions. In addition to that, many species are widespread, which highlights this dispersal capacity. We believe that the widespread distribution of some taxa, especially C. amazonicus and C. longespinicornis, that occur in all biogeographic dominions, might have clouded the biogeographical results. Thus, it is understandable that the ancestral area of the genus might be hard to determine.

Although there are no divergence time estimates for the lineages, it is reasonable to assume that the diversification of eastern South American open vegetation biomes (Caatinga, Cerrado and Chaco), mostly located at the Chacoan dominion, may have played a role in Coccoderus diversification. Some possible vicariant events took place in this region since the Miocene (~15 Ma), and might have shaped the separation of Clade A and B (Werneck 2011): the great Miocene marine transgression that isolated the Central Brazilian Plateaus (Miocene, ~15 MYA); the final uplift of the Central Brazilian Plateau (Late Miocene-Early Pliocene transition, ~5 MYA); the uplift of the Brazilian Plateau along the Espinhaço Range, Serra do Mar and Mantiqueira (Late Pliocene-Early Pleistocene, 2–3 millions years ago); and the subsidence of the Chaco and Pantanal due to the Andean uplift (Late Pliocene-Early Pleistocene, 2–3 millions years ago) (Colli 2005, Porzecanski and Cracraft 2005). Moreover, pleistocene shifts of vegetation that created humid forest corridors, postulated as probable ancient links between Amazon and Atlantic Rainforest and barriers between Caatinga and Cerrado+Chaco (Costa 2003; Sobral-Souza 2015), might have segregated populations and boosted species diversification in the genus, as well as facilitated species with high dispersal capacity to expand their distributional range and establish themselves in other areas.

It is also interesting to point out that only two species have reached Central America, namely C. amazonicus and C. timbara. The closure of the Panama Isthmus (Pliocene, ~3 MYA) (O’Dea et al. 2016) was probably essential for these dispersal events. However, since we do not have a hypothesis for the divergence time of these lineages, any cause for their speciation postulated here is merely speculative.

 5. Conclusion

With the description of Coccoderus trimaculatussp. nov., the species richness of the genus has increased to 15. This study showed the monophyly of Coccoderus with different topologies in the MP, ML, and BI analyses. Additionally, it showed that the Chacoan Dominion (area E) played an important role in the diversification of the genus, and the high dispersal capacity of the species and the overlap of areas made it difficult to determine a single ancestral area. Future studies using molecular data, with larger sampling and fossil calibrations, gave more insights into the biogeographical events in the history of this group.

 6. Declarations

Authors’ contributions. Gabriel S. Ferreira (Conceptualization, investigation, methodology, project administration, resources, supervision, validation, visualization, writing— original draft). André S. Roza (Conceptualization, investigation, methodology, project administration, resources, supervision, validation, visualization, writing— original draft). José Ricardo M. Mermudes (Conceptualized the study, funding acquisition, investigation, resources, writing—review and editing). All authors have read and agreed to the published version of the manuscript.

Competing interests. The authors declare no competing interests.

Ethical aspects. Not applicable

 7. Acknowledgements

We thank Torsten Dikow (curator at USNM) and Charyn Micheli (Museum specialist at USNM). We are grateful to Felipe Barbosa for help in some analyses. We thank to the reviewers for their suggestions for improving the paper. GSF was financed by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior—Brasil (CAPES)—Finance Code 001 (n° 88887.661071/2022-00) and received a grant from the Smithsonian Institution (Ten-Week Graduate Student Fellowship). ASR was financed by the Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) for the personal funding (proc. 205.818/2022 and 205.819/2022). J. R. M. Mermudes is a researcher of the Instituto Nacional de Coleoptera (INCol), a National Institute of Science and Technology (INCT) funded by Brazil’s National Council for Scientific and Technological Development (CNPq grant number 408430/2024-9). He also received funding from CNPq (06105/2016-0, 311679/2019-6, 312786/2022-0) and grants from Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ SEI-260003/006248/2024).

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Files S1–S6

: .zip

File S1. A model selection ran with ModelFinder in IQTree2 [.log file]. – File S2. Morphological characters matrix [.pdf file]. – File S3. Robinson-Foulds distances test [.log file]. – File S4. Maximum Likelihood analysis [.tif file]. – File S5. Bayesian Inference analysis [.tif file]. – File S6. All ancestral area probabilities, as well as recovered possible dispersal and vicariant events [.txt file].

 https://binary.pensoft.net/file/1555960 This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Ferreira GS, Roza AS, Mermudes JRM (2026)