A phylogenetic analysis based on morphology reveals the placement of Lycomesus Zaragoza-Caballero & González-Ramírez, 2019 and Lyconotus Green, 1949 within Lycini (Coleoptera: Lycidae), with the description of a new genus from North America

202 European Journal of Taxonomy 1022: 202–242 https://doi.org/10.5852/ejt.2025.1022.3089 europeanjournaloftaxonomy.eu ISSN 2118-9773 2025 · González-Ramírez M. et al. This work is licensed under a Creative Commons Attribution License (CC BY 4.0) Received: 6 June 2024 • Accepted: 10 July 2025 • Published: 13 October 2025 Topic editor: Tony Robillard • Section editor: Maxwell Barclay • Desk editor: Pepe Fernández Research article urn:lsid:zoobank.org:pub:C59EC983-051A-42BD-A0B6-01DEBAC2F607 A phylogenetic analysis based on morphology reveals the placement of Lycomesus Zaragoza-Caballero & González-Ramírez, 2019 and Lyconotus Green, 1949 within Lycini (Coleoptera: Lycidae), with the description of a new genus from North America Mireya GONZÁLEZ-RAMÍREZ 1,* , Santiago ZARAGOZA-CABALLERO 2 , Juan J. MORRONE 3 & Helga OCHOTERENA-BOOTH 4 1 Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México, 04510 Mexico City, Mexico. 1,2 Laboratorio de Entomología, Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México, 04510 Mexico City, Mexico. 3 Museo de Zoología “Alfonso L. Herrera”, Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México, 04510 Mexico City, Mexico. 4 Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, 04510 Mexico City, Mexico. * Corresponding author: [email protected] 2 Email: [email protected] 3 Email: [email protected] 4 Email: [email protected] Abstract. Lycini are a species-rich tribe within the Lycinae, which are distributed across all biogeographic regions worldwide. Current classification of Lycini has been based exclusively on molecular evidence, and the North American species have been poorly represented. The placement of Lycomesus Zaragoza-Caballero & González-Ramírez, 2019 and Lyconotus Green, 1949 remains uncertain. In order to elucidate the phylogenetic position of these genera within the North American genera and to propose morphological characters that support them, we conducted phylogenetic analyses using parsimony and model-based approaches. Our morphology-based dataset included 91 adult characters from representatives of five genera and 33 species of the Lycini genera of the New and Old Worlds. We propose a new classification of the North American Lycini, consistent with the phylogenetic hypotheses. This includes the following taxa: Lycomesus Zaragoza-Caballero & González-Ramírez, 2019, Lyconotus Green, 1949 stat. rev., Lycorectus González-Ramírez & Zaragoza-Caballero gen. nov., Neolycus Bourgeois, 1883 and Rhyncheros LeConte, 1881. Additionally, we provide a key and diagnoses for all North American Lycini genera, as well as the morphological characters of males and females that define these genera. Our results indicate the necessity for a phylogenetic analysis that incorporates South American Lycini. This will enable a more precise generic classification of American Lycini. GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 203 Keywords. Net-winged beetle, Lycinae, taxonomy, Nearctic region, Neotropical region. González-Ramírez M., Zaragoza-Caballero S., Morrone J.J. & Ochoterena-Booth J. 2025. A phylogenetic analysis based on morphology reveals the placement of Lycomesus Zaragoza-Caballero & González-Ramírez, 2019 and Lyconotus Green, 1949 within Lycini (Coleoptera: Lycidae), with the description of a new genus from North America. European Journal of Taxonomy 1022: 202–242. https://doi.org/10.5852/ejt.2025.1022.3089 Introduction Lycini Laporte, 1836 are the most diverse tribe within the subfamily Lycinae Laporte, 1836, with eight genera and more than 400 described species (Kusy et al. 2020). Lycines can be distinguished from other lycids by the elongated head forming a ‘rostrum’; slender mandibles; antennomeres 3–10 flat, serrate to parallel-sided; elytra slightly to distinctly broadened posteriorly; spoon-shaped phallobase; and long and slender phallus (LeConte 1881; Green 1949; Zaragoza-Caballero 1995; Bocak & Bocakova 2008; Kusy et al. 2019, 2020). The tribe has a wide distribution, in almost all biogeographical regions of the world except the Antarctic (Gorham 1880; Kleine 1933; Blackwelder 1945; Green 1949; Bocak & Bocakova 2008; Masek et al. 2018). It has been proposed that they originally evolved in the southern part of the Nearctic region and Mesoamerica and later dispersed to Eastern Asia (Masek et al. 2018; Kusy et al. 2020). Despite their hypothesized American origin, approximately 70% of species are found in the Afrotropical region (Masek et al. 2018; Kusy et al. 2020). In contrast, only 7.6% of their diversity is reported in the Nearctic and Neotropical regions, with only four genera represented (Celiasis Laporte, 1840, Rhyncheros LeConte, 1881, Neolycus Bourgeois, 1883 and Lycomesus Zaragoza-Caballero & González-Ramírez, 2019) and 45 species reported for the region (Chevrolat 1834; Gorham 1880; Kleine 1933; Blackwelder 1945; Green 1949; Zaragoza-Caballero 1995; Pérez-Hernández et al. 2019; ZaragozaCaballero & González-Ramírez 2019; Kusy et al. 2020). The genus Celiasis is reported in the literature to be endemic to Colombia (Gorham 1880; Kleine 1933; Blackwelder 1945). Since the type species could not be found, previous researchers dealing with this taxon (Kusy et al. 2020) could not study it, and this taxon is now considered a nomen dubium (Evenhuis 2012; Kusy et al. 2020). Bourgeois (1906), however, proposed that Celiasis mirabilis Lacordaire, 1857 (the type species of Celiasis) is not a lycid, but rather belongs to a genus of lycidiform melyrids that was later described by Blanchard under the name Chalcas Blanchard, 1845 (Ch. trabeatus Fairmaire, 1847). Despite the compelling story and rationale by Bourgeois (1906), Celiasis remain a nomen dubium in Lycini until the South American lineages are treated in the future. Furthermore, studies addressing the taxonomy of American genera have been limited. Consequently, authors often offer only cursory discussions on the morphology of the taxa studied (Chevrolat 1834; Melsheimer 1846; Dugès 1878; Gorham 1880, 1884; LeConte 1881; Green 1949). Little informative descriptions, ambiguous taxonomic delimitations of genera and subgenera without a phylogenetic context, and inaccessibility of type material (Bourgeois 1883; Pic 1922; Kusy et al. 2020) have hindered the systematic work on the Lycini fauna worldwide (Kusy et al. 2020). In the New World, research on Lycini has been limited, with the majority of work focusing on catalogues, new species descriptions, checklists, and immature stages descriptions (Chevrolat 1834; Gorham 1880; Dugès 1896; Kleine 1933; Blackwelder 1945; Green 1949; Zaragoza-Caballero & González-Ramírez 2019; Pérez-Hernández et al. 2019; González-Ramírez & Zaragoza-Caballero 2024). Recently, Kusy et al. (2020) proposed a new genus-level arrangement of lycines based on a molecular phylogenetic analysis that included approximately 100 species. Their classification includes seven genera (Celiasis, Neolycus, Rhyncheros, Lipermes Waterhouse, 1879, Haplolycus Bourgeois, 1883, Lycostomus Motschulsky, 1861 and Lycus Fabricius, 1787). Kusy et al. (2020) separated the Old and New World species and treated Lyconotus Green, 1949 as a synonym of Rhyncheros. Despite the important contributions in this work, European Journal of Taxonomy 1022: 202–242 (2025) 204 the North American species of Lycini were underrepresented in their study, and the generic assignment of Lycomesus and Lyconotus raised several questions. In order to contribute to the knowledge of the tribe and to evaluate the phylogenetic relationships of the North American Lycini, we conducted a phylogenetic analysis with dense taxon sampling aimed to encompass the largest number of genera and species from North and South America. Our objectives are to test the monophyly of Lycini, to clarify the phylogenetic relationships of North American genera, and to propose a classification consistent with the phylogenetic hypothesis. Finally, we utilize the results obtained to define the diagnostic characters of North American genera of Lycini, with which a dichotomous key has been constructed. Additionally, a diagnosis of the genera is presented. Material and methods Taxon sampling Forty species were included in the phylogenetic analysis (Table 1). The ingroup consisted of 29 nominal species of Lycini and four undescribed species belonging to five genera (Lycomesus, Lycostomus, Lycus, Neolycus and Rhyncheros). The outgroup was represented by six species of the tribe Calopterini Green, 1949, which is considered to be more closely related to Lycini (Bocak & Matsuda 2003). For rooting the cladogram, we used a more distantly related species, Chauliognathus sp. (Cantharidae). Repositories A total of 1060 specimens (Supp. file 1) from the following collections (curators in parentheses) were analyzed. ANSP = Academy of Natural Sciences, Philadelphia, USA (Jon Gelhaus) CAS = California Academy of Sciences, California, San Francisco, USA (Chris Grinter) CNIN = Colección Nacional de Insectos, Instituto de Biología, UNAM, Mexico City (Santiago Zaragoza Caballero) NHMUK = Natural History Museum, London, UK (Michael Geiser) Due to the lack of suitable material, the character coding for the male of Lyconotus lateralis (Melsheimer, 1846) was based on information obtained from the original description (Melsheimer 1846) and Green (1949); the coding of the female of Lycus trabeatus Guérin-Méneville, 1835, Rhyncheros godmani (Gorham, 1880), Rhyncheros nigrofumosus (Hinton, 1933), and the male of Neolycus carmelitus (Gorham, 1880) was based on photographs of the type specimens held at the NHMUK and CAS; the coding of the male of Lycostomus praeustus Fabricius, 1792 was based on the information provided by Kazantsev (2003); and the coding for the female of Rhyncheros sanguinipennis Say, 1823 was based on photographs of specimens from the ANSP (the acronyms CNIN, and ANSP follow the Insect and Spider Collections of the World website [Evenhuis 2023]). Characters All examined specimens were studied using a Zeiss Discovery V8 stereo microscope equipped with an ocular micrometer that was used for measurements. Genitalia were dissected and the surplus tissue was macerated using 10% KOH for 5 minutes. The wing morphology nomenclature and interpretation are based on Lawrence et al. (2021), and we followed Kazantsev (2003) for the female genitalia descriptions. Due to the complexity to appropriately codify character states of characters 70 (paramere shape in lateral view) and 72 (phallus shape), we conducted a geometrical morphometric analysis using a representative ‘normal’ sample and one image per species, all using the same position and scale (Vega-Badillo et al. 2021). Photographs were captured with a Zeiss Axio Zoom V16 microscope with an Axiocam GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 205 Table 1 (continued on next page). Taxon sampling, with indication of the country. Family Tribe Species Country Ingroup Lycidae Lycini Lycomesus llorentei Zaragoza-Caballero & González-Ramírez, 2019 Mexico Lycus ampliatus (Fåhraeus, 1851) South Africa Lycus trabeatus Guérin-Méneville, 1835 Burkina Faso Lycostomus praeustus Fabricius, 1792 India Lycostomus rubrocinctus Fairmaire, 1886 China Neolycus arizonensis (Green, 1949) Mexico and USA Neolycus carmelitus (Gorham, 1880) Mexico Neolycus fernandezi (Dugès, 1878) Mexico and USA Neolycus lecontei (Green, 1949) Mexico Neolycus sallaei (Gorham, 1880) Mexico Neolycus schönherri (Chevrolat, 1834) Mexico Neolycus scutellatus (Gorham, 1880) Mexico Rhyncheros sp. 1 Mexico Rhyncheros sp. 2 Mexico Rhyncheros sp. 3 Mexico Rhyncheros sp. 4 Mexico Rhyncheros carnifex (Gorham, 1880) Mexico Rhyncheros fuliginosus (Gorham, 1880) Mexico Rhyncheros fulvellus (LeConte, 1881) Mexico and USA Rhyncheros fulvellus femoratus (LeConte, 1881) Mexico Rhyncheros godmani (Gorham, 1880) Mexico Rhyncheros lateralis (Melsheimer, 1846) USA Rhyncheros lineicollis (Chevrolat, 1834) Mexico Rhyncheros loripes (Chevrolat, 1834) Mexico Rhyncheros minutus (Green, 1949) Mexico and USA Rhyncheros nigrofumosus (Hinton, 1933) Mexico Rhyncheros rusticus (Gorham, 1884) Mexico Rhyncheros sagittatus (Green, 1949) Mexico Rhyncheros sanguineus (Gorham, 1884) Mexico Rhyncheros sanguinipennis Say, 1823 Mexico Rhyncheros semiustus (Chevrolat, 1834) Mexico Rhyncheros simulans (Schaeffer, 1911) Mexico Rhyncheros sordidus Gorham, 1880 Mexico European Journal of Taxonomy 1022: 202–242 (2025) 206 Table 1 (continued). Taxon sampling, with indication of the country. Family Tribe Species Country Outgroup Lycidae Calopterini Calopteron bifasciatum Gorham, 1884 Mexico Calopteron jimenezi Dugés, 1878 Mexico Calopteron corrugatum Candeze, 1861 Mexico Calopteron discrepans Newman, 1838 Mexico Caenia sinuata Kirsch, 1865 Mexico Caenia amplicornis LeConte, 1881 USA Cantharidae Chauliognathus sp. Mexico Fig. 1. Designation of landmarks and semilandmarks for the geometric morphometric analysis. A. In character 70, three landmarks were selected in the left paramere (one on the right side, one on the apex, and one on the left side), and 40 semilandmarks. B. In character 72, three landmarks were selected in the phallus (one on the right, another on the apex, and a third on the left), and 70 semilandmarks. GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 207 MRC5 digital camera and Lens Plan NeoFluar, at the Laboratorio de Microscopía y Fotografía de la Biodiversidad at the Instituto de Biología, UNAM ( Universidad Nacional Autónoma de México) . We registered landmark (LM) and semilandmark (SLM) points with each digital photograph using tpsDig2 ver. 2.3.2 (Rohlf 2017). For character 70, we selected three landmarks on the left paramere: one on the right side, one on the apex, and one on the left side. The landmarks on the right and left sides were situated at the point where the paramere joins the phallobase, and the posterior landmark was placed at the apex of the paramere (3 LM, and 40 SLMs) (Fig. 1A). For character 72, we selected three landmarks on the phallus: one on the right, another on the apex, and a third on the left. The landmarks on both sides were located at the junction between the phallus and the parameres, while the posterior landmark was located at the apex of the phallus (3 LM, and 70 SLMs) (Fig. 1B). All outlines were registered as SLMs using tpsDig2 ver. 2.3.2 (Rohlf 2017), consistently registering equivalent points along outlines ‘by length’. To identify the groups with the highest similarity in shape for these structures (character states), we conducted a hierarchical cluster analysis for each structure using an Euclidean distance matrix. The difference matrix obtained was plotted on a dendrogram, which served as a guide for the character state assignment (Figs 2–3, respectively). The optimal number of clusters was determined using the ‘elbow’ method. All cluster analyses were conducted using RStudio software (R Core Team 2021) with ggplot2 (Wickham 2016) and factoextra (Kassambara & Mundt 2020). Many characters included in this study were previously used in phylogenetic analyses of the family (Kazantsev 2003, 2013; Bocakova 2005) or for species descriptions (Chevrolat 1834; Gorham 1880; Green 1949; Zaragoza-Caballero 1995; Zaragoza-Caballero & González-Ramírez 2019). Several characters are used here for the first time and are indicated by an asterisk in the character list (see below). A total of 91 characters were derived from the external morphology of adult specimens, 47 of which are scored here for the first time for all included taxa. Twenty characters were obtained from male and female genitalia, 15 of which are reported here for the first time for all included taxa. A total of 31 binary and 60 multistate characters were coded, three characters (4, 27 and 42) were coded as polymorphic for at least some terminals. For continuous characters (1, 28, 41, 42, 52, 77, 80, 84), the designation of character states was carried out in accordance with the formula proposed by Vega-Badillo et al. (2021). This entailed calculating the difference between the minimum and maximum values (dmm) and dividing it by the number of character states. The resulting value was then expressed as dmm = max – min / 3. Subsequently, character states were assigned as follows: 0 = min + dmm; 2 = max – dmm; and 1 = intervals between 0 and 2. Individual consistency and retention indices (CI, RI) are provided for all characters from the implied weights analysis (k = 7.187500; see below under Phylogenetic analysis). Character and character states Head 0. Coronal sulci: deeply divided (0) (Fig. 4A); slightly divided (1) (CI = 0.50; RI = 0.88). 1. *Width of interantennal distance: less than 0.09 mm (0) (Fig. 4B); 0.1 mm–0.12 mm (1) (Fig. 4C); more than 0.13 mm (2) (Fig. 4D) (CI = 0.40; RI = 0.76). 2. Male IV–X antennomeres shape: bidentate (0) (Fig. 4E); unidentate (1) (Fig. 4F); rectangular (2) (Fig. 4G); filiform (3) (CI = 0.30; RI = 0.58). 3. Female IV–X antennomeres shape: bidentate (0); unidentate (1); rectangular (2); filiform (3) (CI = 0.33; RI = 0.40). 4. *First three antennomeres colour: black (0); yellow (1); bicolour (2) (CI = 0.40; RI = 0.75). European Journal of Taxonomy 1022: 202–242 (2025) 208 5. Antennomere III proportional length: elongate, almost subequal in length to antennomere IV (0); short, about 0.3 mm shorter than antennomere IV (1); elongate, about 0.7 mm longer than antennomere IV (2) (CI = 0.66; RI = 0.91). 6. Head with rostrum: absent (0); present (1) (CI = 1; RI = 1). 7. *Rostrum relative length: wider than long (0) (Fig. 4H); as long as wide (1) (Fig. 4I); longer than wide (2) (Fig. 4J) (CI = 0.40; RI = 0.50). Fig. 2. Dendrogram obtained after digitizing the paramere contour using Euclidian distances, showing the six recognized characters states (character 70). GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 209 8. *Rostrum length-width ratio: less than 1.9 (0); 2 to 2.6 (1); more than 2.7 (2) (CI = 0.22; RI = 0.22). 9. *Labrum shape in dorsal view: oval (0) (Fig. 4K); semi-circular (1); semi-square (2) (Fig. 4L); rectangular (3) (Fig. 4M); obcordate (4); bilobate (5) (CI = 0.50; RI = 0.70). 10. *Labrum width-length relation: as wide as long (0); wider than long (1); longer than wide (2) (CI = 0.14; RI = 0.45). Fig. 3. Dendrogram obtained after digitizing the phallus contour using Euclidian distances, showing the six recognized characters states (character 72). European Journal of Taxonomy 1022: 202–242 (2025) 210 Fig. 4. Morphology of Lycini. A. Frontal hump of Lyconotus sp. 2, ♂ (CNIN, QR 70966). B–D. Interantennal distance. B. Neolycus fernandezi (Dugès, 1878), ♂ (CNIN, QR 67699). C. Lyconotus sp. 1, ♂ (CNIN, QR 70971). D. Rhyncheros minutus (Green, 1949), ♀ (CNIN, QR 68274). E–G. Antenna. E. Neolycus scutellatus (Gorham, 1880), ♂ (CNIN, QR 67865). F. Neolycus fernandezi, ♂ (CNIN, QR 67714). G. Lyconotus semiustus (Chevrolat, 1834), ♂ (CNIN, QR 68724). H–J. Rostrum. H. Rhyncheros carnifex (Gorham, 1880), ♀ (CNIN, QR 67878). I. Rhyncheros lineicollis (Chevrolat, 1834), ♂ (CNIN, QR 67905). J. Rhyncheros loripes (Chevrolat, 1834), ♀ (CNIN, QR 68185). K–M. Labrum. K. Lyconotus sp. 3, ♀ (CNIN, QR 70975). L. Rhyncheros minutus, ♂ (CNIN, QR 68294). M. Neolycus scutellatus, ♂ (CNIN, QR 67856). N–P. Maxillary palpomere. N. Neolycus schoenherri (Chevrolat, 1834), ♂ (CNIN, QR 67841). O. Lycostomus rubrocinctus Fairmaire, 1886, ♂ (CNIN, China). P. Neolycus arizonensis (Green, 1949), ♂ (CNIN, QR 67730). Q–X. Pronotum. Q. Lycostomus rubrocinctus, ♂ (CNIN, China). R. Neolycus arizonensis, ♂ (CNIN, QR 67734). S. Neolycus schoenherri, ♂ (CNIN, QR 67820). T. Rhyncheros loripes, ♂ (CNIN, QR 68093). U. Rhyncheros lineicollis, ♂ (CNIN, QR 67913). V. Lyconotus sp. 2, ♂ (CNIN, QR 70966). W. Rhyncheros fuliginosus (Gorham, 1880), ♂ (CNIN, QR 67757). X. Lyconotus sp. 4, ♂ (CNIN, QR 70967). GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 217 70. *Parameres shape in lateral view was assigned to a group through of geometrical morphometric analysis (Fig. 2): group 0 (0); group 1 (1); group 2 (2); group 3 (3); group 4 (4); group 5 (5) (CI = 0.38; RI = 0.27). 71. Parameres apex shape: rounded (0) (Fig. 7E); acute (1) (Fig. 7F); acuminate (2) (Fig. 7G) (CI = 0.18; RI = 0.52). 72. *Phallus shape in lateral view was assigned to a group through of geometrical morphometric analysis (Fig. 3): group 0 (0); group 1 (1); group 2 (2); group 3 (3); group 4 (4); group 5 (5) (CI = 0.50; RI =0.70). 73. Pair of thorns on phallus: absent (0) (Fig. 7H); present (1) (Fig. 7I) (CI = 0.25; RI = 0.76). 74. *Phallus thorns location: in ventral view (0) (Fig. 7J); in lateral view (1) (Fig. 7K) (CI = 0.50; RI = 0.83). 75. *Phallus thorns relative position: near parameres (0); around phallus mid-length (1) (CI = 0.33; RI = 0.71). 76. *Phallus thorns modified to receive parameres: absent (0); present (1) (CI = 1; RI = 1). 77. *Phallus thorns relative length: less than 0.046 mm (0); 0.047 mm–0.083 mm (1); more than 0.084 mm (2). (CI = 0.28 RI = 0.16). 78. *Phallus thorns apex shape: truncated (0); rounded (1); acute (2); acuminate (3). (CI = 0.50; RI = 0.66). 79. *Phallus apex shape: rounded (0); sharp (1); bifurcated (2); pincer (3) (CI = 1; RI = 1). Female genitalia 80. *Stylus length: less than 0.21 mm (0); 0.22 mm–0.29 mm (1); more than 0.30 mm (2) (CI = 0.25; RI = 0.40). 81. *Stylus shape: robust (0); slender (1) (CI = 0.11; RI = 0.20). 82. *Outer margin of the stylus cleft: absent (0); present (1) (CI = 1; RI = 1). 83. *Coxites-valvifers integration: non-joined and separated (0) (Fig. 7L); joined (1); merged (2) (Fig. 7M); non-joined and overlapping structures (3) (Fig. 7N) (CI = 0.75; RI = 0.94). 84. *Coxites length: less than 0.97 mm (0); 0.98 mm–1.50 mm (1); more than 1.51 mm (2) (CI = 0.33; RI = 0.55). 85. Coxites shape: almost straight (0); emarginate medially (1); emarginate apically, S-shaped (2) (CI = 0.22; RI = 0.46). 86. *Outer margin of the coxites cleft: absent (0); present (1) (CI = 1; RI = 1). 87. *Coxites inner apical margins relative position: nearby (0); distant (1) (CI = 0.33; RI = 0.81). 88. *Length relation between coxites and valvifers: shorter (0); coxites as long as valvifers (1); longer (2) (CI = 50; RI = 0.85). European Journal of Taxonomy 1022: 202–242 (2025) 218 89. Fusion of valvifers at the base: absent (0); present (1) (CI = 0.50; RI = 0.90). 90. *Coxites inner margin base shape: concave (0); lobed (1) (CI = 1; RI = 1). Phylogenetic analysis The matrix was created using Winclada ver. 1.00.08 (Nixon 2002). Multistate characters were treated as non-additive. Unknown character states were coded with question marks (‘?’), which was mostly the case when only one sex was examined due to the lack of specimens for the other sex; inapplicable character states were coded using a dash (‘-’) when the preceding character in the matrix was coded as absent. Total polymorphism (all possible character states present in the same taxon) is represented by WinClada using an asterisk (‘*’); partial polymorphisms (only some of the possible character states present in the same taxon) are represented by WinClada using dollar symbols (‘$’). Phylogenetic analyses were carried out using parsimony, maximum likelihood (ML), and Bayesian inference (BI). Parsimony analyses were conducted using TNT ver. 1.6 (Goloboff & Morales 2023) under equal weights (EW) and heuristic searches used the ratchet method (Nixon 1999). The specific parameters used were: 1000 random seeds, find minimum length at least 10 times, and 5000 iterations. The analysis was conducted on multiple occasions until we established that the results were stable. Winclada ver. 1.00.08 (Nixon 2002) was used to construct a strict consensus tree (Nixon & Carpenter 1996) from the most parsimonious trees. These trees were evaluated by tree length (L), consistency index (CI), and retention index (RI). To examine the effect of individual character homoplasy on the analyses results, we conducted a second analysis using implied weights (IW; Goloboff 1993). We used the TNT script (setk.run) written by Salvador Arias (Hermes et al. 2014) to determine the optimal value for the constant k (Goloboff et al. 2008). A value of 7.187500 was returned as the most appropriate k value and was therefore used in all subsequent implied weighting schemes employing the heuristic method of new search strategies (ratchet), with the following parameters: 1000 random seeds, find minimum length 10 times and 5000 iterations. The optimal trees were opened in Winclada ver. 1.00.08 (Nixon 2002) to calculate the length and consistency and retention indices as well as to construct their strict consensus tree (Nixon & Carpenter 1996). To assess the statistical branch support, a bootstrap (BS) and a jackknife (JS) analyses were carried out with NONA (Goloboff 1994) through WinClada ver. 1.00.08 (Nixon 2002). Ten thousand replicates were conducted utilizing 10 initial trees, holding 10 trees and expanding the memory up to 1000 trees (mult*10 hold/10 max*10 000). We considered strong support above 80%. The character optimizations were mapped on the strict consensus tree in Winclada ver. 1.00.08 (Nixon 2002). Three different optimizations were employed: unambiguous changes, and accelerated (ACCTRAN) and delayed (DELTRAN) transformations (Supp. file 4: figs S1–S3). The decision to employ DELTRAN was motivated by the fact that, in certain instances, when there are terminals with unknown or inapplicable states, ACCTRAN considers it to be a spurious synapomorphy. In contrast, DELTRAN does not execute this transformation, considering an apomorphy exclusively for taxa that possess the given state (De Santis & Nihei 2022). The presence of artefacts in the mapping process was investigated through a thorough examination of polytomies. This involved a meticulous comparison of the location of character states among the most parsimonious or optimal trees in branches supporting polytomies. Toggling the options to map characters vs character states was used to verify the distribution of character states among our taxon sample for identifying diagnostic characters. IQ-Tree 2 software (Minh et al. 2020) was used for maximum likelihood inference (ML). The evolutionary model was obtained using ModelFinder (Kalyaanamoorthy et al. 2017) implemented in IQ-Tree 2. According to the Bayesian information criterion (BIC), the best fit model was the k-states Markov Mkv (MK) (Lewis 2001) with verification bias correction (ASC), gamma distribution using four categories of discrete rates (G4), and equal state frequencies (FQ). Branch support was estimated by ultrafast bootstrapping (UFBoot) using 10 000 replicates (Hoang et al. 2018). Values ≥ 95% were considered indicative of strong support. GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 219 Bayesian inference (BI) was performed using Mr. Bayes ver. 3.2.7 (Ronquist et al. 2012), using the Mkv model (Ronquist & Huelsenbeck 2003) for variable morphological characters, with gamma for state frequencies and 0.07 for temperature. Four simultaneous runs of two million generations were run, each with one cold chain and three heated chains. Samples were drawn every 500 Markov chain Monte Carlo steps, with the first 25% discarded as burn-in. The run was automatically stopped when the average standard deviation of split frequencies was below 0.01. The posterior probability (PP) was employed to provide support for the nodes (Yang & Rannala 1997), values ≥ 95% were interpreted as high nodal support and ≥ 80% for standard BS values. Phylogenetic trees were visualized in FigTree ver. 1.4.4 (Rambaut 2009). The four phylogenetic hypotheses were subjected to statistical analysis to determine their statistical significance (p-value of 0.05). To this end, the following parsimony-based tree topology tests were conducted: Templeton test and winning-sites test, implemented in PAUP ver. 4.0a software (Swofford & Bell 2017), and likelihood-based tests: the weighted Kishino-Hasegawa (KH) test, the weighted Shimodaira-Hasegawa (SH) test and Shimodaira’s approximately unbiased (AU) test with IQ-Tree 2 software (Minh et al. 2020). Results Phylogenetic analysis Input data The data matrix comprises 219 cells coded as missing (?), 176 as not applicable (-), five as full polymorphisms, and 23 as partial polymorphisms. The complete matrix used for the morphological analysis can be found in Supp. file 2: table 1. The application of different approaches yielded congruent patterns on major linages. Parsimony analysis using equal weights The parsimony analysis under EW yielded 4 most parsimonious trees, with a length of 581 steps, a consistency index (CI) of 0.32, and a retention index (RI) of 0.63. The strict consensus tree has 587 steps (CI = 0.31; RI = 0.63) (Supp. file 3: fig. S1). The tribe Lycini was recovered as monophyletic (BS = 70; JS = 79), but Lycostomus, Rhyncheros and Neolycus were not recovered as monophyletic. However, three clades within the tribe were recovered as monophyletic: Lycus, Lycomesus, and Lyconotus (BS = 85; JS = 89) (Supp. file 3: fig. S1). Parsimony analysis using implied weights The parsimony analyses under IW with a constant k = 7.187500 using the TNT script (setk.run) resulted in one most parsimonious tree (L = 583, CI = 0.31, RI = 0.63) (Supp. file 4: fig. S2). Bootstrap and jackknife values for each branch are shown in Fig. 8. As is common with morphological matrices, most clades received low support (Bocakova 2001, 2005; Nascimento et al. 2020; Vega-Badillo et al. 2021). The cladogram under IW shows that the Lycini are monophyletic (BS = 69; JS = 80), and their sister group is Caenia + Calopteron (Fig. 8). The monospecific genus Lycomesus was recovered as the sister group to Lyconotus (Fig. 8), and Lyconotus was recovered as monophyletic (BS = 84; JS = 88). However, Lycostomus was not found to be monophyletic (Fig. 8). Lycus was recovered as sister to the remaining North American Lycini (Fig. 8). Lycorectus sordidus (Gorham, 1880) gen. et comb. nov. was recovered as sister to Neolycus and the rest of Rhyncheros (Fig. 8). The analysis shows that Neolycus was monophyletic (BS = 59; JS = 64). Fourteen species of Rhyncheros were forming a clade (Fig. 8). Maximum likelihood analysis The ML analysis yielded a topology that was well resolved, exhibiting a high degree of congruence with the topology obtained under IW (Supp. file 5: fig. S1). The monophyly of the Lycini was recovered with European Journal of Taxonomy 1022: 202–242 (2025) 220 strong nodal support (UFBoot = 99%). The analysis revealed that the clade Lycomesus + Lyconotus was a sister group to all Lycini. Lycostomus was recovered as non-monophyletic. The monophyly of Lyconotus and Lycus was well supported with 96% and 91% of UFBoot, respectively. The analysis further revealed Rhyncheros to be paraphyletic, Lycorectus sordidus gen. et comb. nov. was recovered as sister group to Neolycus, and the monophyly of Neolycus was supported by a low UFBoot value (80%) (Supp. file 5: fig. S1). Fig. 8. A single cladogram of parsimony analysis under implied weights (L = 583 CI = 0.31; RI = 0.63). above branches indicate bootstrap values and numbers below the branches corresponds to the jackknife values (> 50). GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 221 Bayesian inference Conversely, the cladogram obtained by BI (Supp. file 6: fig. S1) exhibited a lower resolution than those obtained by parsimony under EW, IW and ML. The cladogram shows that the Lycini are monophyletic (PP = 0.97), and Rhyncheros was recovered as polyphyletic, a clade comprising Rhyncheros [R. fuliginosus (Gorham, 1880), R. nigrofumosus (Hinton, 1933), R. carnifex (Gorham, 1880) and R. godmani] (PP = 0.28) was recovered as early diverging, in a sister position to other Lycini. A second clade of Rhyncheros [R. lineicollis (Chevrolat, 1834) and R. sanguinipennis] (PP = 0.42) was identified as a sister group to remaining members of the Lycini. The clade Neolycus scutellatus (Gorham, 1880) + Rhyncheros sanguineus (Gorham, 1884) was identified as a sister group to a third Rhyncheros clade [R. rusticus (Gorham, 1884), R. fulvellus (LeConte, 1881), R. fulvellus femoratus (LeConte, 1881), R. simulans (Schaeffer, 1911), R. loripes (Chevrolat, 1834), R. sagittatus (Green, 1949) and R. minutus (Green, 1949)] (PP = 0.59). Lycorectus sordidus gen. et comb. nov. was found in a clade with the other species of Neolycus, and Neolycus was recovered as a paraphyletic group. The genus Lycus was recovered as monophyletic (PP = 0.83), whereas Lycostomus was recovered as paraphyletic. Furthermore, Lycomesus was identified as the sister group to Lyconotus, and the monophyly of Lyconotus was well supported (PP = 0.94). The findings of the implementation of the tree topology tests (Table 2) demonstrated that, among the parsimony-based tree topology tests, the trees generated by parsimony under EW, IW and ML (Table 2; Fig. 8; Supp. file 3: fig. S1, Supp. file 5: fig. S1) outperformed the BI reconstructions (Templeton test, p > 0.210; winning-sites test, p < 0.092) (Table 2; Supp. file 6: fig. S1). Similarly, among the likelihoodbased tree topology tests, the trees generated by parsimony under IW and ML (Table 2; Fig. 8; Supp. file 5: fig. S1) performed better than the parsimony under EW and BI (Table 2; Supp. file 3: fig. S1, Supp. file 6: fig. S1). According to the applied topology tests, trees generated by parsimony under IW (Fig. 8) and ML (Supp. file 5: fig. S1) are not significantly different and are equally plausible (parsimony under IW: Templeton test, best; winning-sites test, best; W-KH, P > 0.466; W-SH, p > 0.718; AU, p > 0.564. ML: Templeton test, p > 0.454; winning-sites test, p > 0.472; W-KH, p > 0.534; W-SH, p > 0.843; AU, p > 0.636). However, in order to maintain the taxonomic stability of the group (Ferreira et al. 2023), the preferred topology is presented, based on parsimony under IW. This topology (Fig. 8) proposes a new classification for North American Lycini, with the genus Lyconotus reinstated (Fig. 8) and a new genus, Lycorectus (Fig. 8), established. Furthermore, a key is provided to facilitate the recognition of the genera of Lycini in North America, based on the characters that support the groups in the cladogram (Fig. 8; Supp. file 4: fig. S2). Table 2. Results of tree topology tests. P-EW signifies parsimony under equal weights, whilst P-IW denotes parsimony under implicated weights, ML indicates maximum likelihood, and BI indicates Bayesian inference for each topology. Parsimony-based tests: Templeton and winning-sites were performed on the PAUP ver. 4.0a software (Swofford & Bell 2017). Likelihood-based topology tests: weighted Kishino-Hasegawa test (W-KH), weighted Shimodaira-Hasegawa test (W-SH), and Shimodaira’s approximately unbiased test (AU) were performed on IQTree 2 software (Minh et al. 2020). Grey color denotes p-value > 0.05 (statical significance). ‘best’ means that the tree was the best tree for a given optimality criterion. Topology Templeton winning-sites W-KH W-SH AU P-EW 0.627 0.486 0.233 0.411 0.240 P-IW best best 0.466 0.718 0.564 ML 0.454 0.472 0.534 0.843 0.636 BI 0.210 0.092 0.294 0.483 0.352 European Journal of Taxonomy 1022: 202–242 (2025) 222 Key to genera of Lycini from North America 1. Length of antennomere III almost the same as antennomere IV (Fig. 4G); head with rostrum longer than wide, but very slender at base; subquadrangular or oval labrum; trochanters with acute internal angle; metatibial spurs short, and acute (Fig. 6F) ............................................................................. 2 – Length of antennomere III longer than antennomere IV (Fig. 4E–F); head with rostrum of variable length, but slightly to strongly widened at base; semi-circular to semi-square labrum; trochanters with blunt internal angle; metatibial spurs long, and apically variable shape (Fig. 6G) .................. 3 2. Labrum with truncated anterior margin; in males, last abdominal sternite and phallus are curved at an angle of 45°, without a pair of thorns in the middle part of phallus, without modifications to receive the parameres, which are rounded (Fig. 7E) ....................................................................................... ..............................................................Lycomesus Zaragoza-Caballero & González-Ramírez, 2019 – Labrum with rounded anterior margin; in males, last abdominal sternite and phallus are curved at an angle of 90° (Fig. 6P), with a pair of thorns in the middle part of phallus, phallus modified to receive parameres, these are triangular (Fig. 7K) ....................................... Lyconotus Green, 1949 stat. rev. 3. Elytra with sexual dimorphism, male considerably dilated in middle part, while female with moderately dilated elytra (Fig. 9D–E); radiomedial loop broadly curved (Fig. 5M); metatibial spurs unequal (inner acute, outer slightly broader apically and bluntly rounded), and situated close together at the base (Fig. 6E) .................................................................................. Neolycus Bourgeois, 1883 – Elytra lacking sexual dimorphism (Figs 9F–G, 10A, 11A); radiomedial loop slightly curved (Fig. 5N); metatibial spurs equal, acute, and with a considerable distance between them at the base (Fig. 6E, G) ....................................................................................................................................... 4 4. Males with concave or emarginate posterior border on the seventh sternite; thorns on the phallus present or absent (Fig. 7C, F, I) ..............................................................Rhyncheros LeConte, 1881 – Males with convex posterior border on the seventh sternite; thorns on the phallus absent (Fig. 11C– E) .................................................... Lycorectus González-Ramírez & Zaragoza-Caballero gen. nov. Taxonomy Class Insecta Linnaeus, 1758 Order Coleoptera Linnaeus, 1758 Superfamily Elateroidea Family Lycidae Laporte, 1836 Subfamily Lycinae Laporte, 1836 Tribe Lycini Laporte, 1836 Genus Lycomesus Zaragoza-Caballero & González-Ramírez, 2019 Figs 7E, 9A Lycomesus Zaragoza-Caballero & González-Ramírez, 2019: 99. Type species Lycomesus llorentei Zaragoza-Caballero & González-Ramírez, 2019. Diagnosis Lycomesus shares similarities with Lyconotus, but it differs in the shape of the anterior margin of the labrum, which is truncated; the presence of a radiomedial loop with sharp edges; the structure of the aedeagus, with parameres that are short and rounded, last abdominal sternite and phallus are characterised GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 223 by an angled curvature measuring 45°, as well as the lack of thorns on the phallus. Additionally, the phallus is not modified for the reception of the parameres (Fig. 7E). Material examined Refer to Supp. file 1. Redescription Body slender. Head concealed by pronotum, rostrum long and slightly widened. Interantennal distance less than 0.09 mm. Antennomere III almost as long as antennomere IV. Labrum semi-square. Mandibles longer than labrum. Pronotum widest at base, with anterior border rounded and anterior margins not prominent, pronotal margin less than 0.52 mm in wide. Elytra with three distinct costae on each elytron (Fig. 9A). Radiomedial loop angle on wing sharp. Trochanters with acute internal angle, metatibial spurs equal and short. Last sternite partially curved. Male genitalia with short, rounded parameres, phallus laterally compressed, slightly widened, and curved at apex (curvature of about 45°), phallus without thorns (Fig. 7E). Distribution Lycomesus has only been reported from the type locality in Mexico (Nearctic region) (ZaragozaCaballero & González-Ramírez 2019). Genus Lyconotus Green, 1949 stat. rev. Figs 4A, C, G, K, V, X, 5D, H, L, O, Q, 6C, F, J, N, P, 7K, N, 9B–C Lyconotus Green, 1949: 67. Type species Lycus lateralis Melsheimer, 1846. Diagnosis Similar in appearance to Lycomesus (Fig. 9B–C), but differs in the oval shape of the labrum (Fig. 4K); the acute apex of the elytra; the membranous wing with the radiomedial loop slightly acute (Fig. 5O); in males the last sternite and phallus curve at 90°, the thorns of the phallus are modified to receive parameres (Fig. 7K). In the female genitalia, the coxites and valvifers are independent structures, and the outer margin of the coxites cleft is present (Fig. 7N). Material examined Refer to Supp. file 1. Redescription Body slender. Head concealed by pronotum, rostrum long and very slender (Fig. 4A). Interantennal distance around 0.1–1.12 mm (Fig. 4C). Antennomere III almost as long as antennomere IV (Fig. 4G). Labrum oval (Fig. 4K). Mandibles longer than labrum. Pronotum widest at base, with anterior border rounded and anterior margins not prominent, between 0.53–0.81 mm in width. Elytra without sexual dimorphism, with three distinct costae on each elytron (Fig. 9B–C). Radiomedial loop angle slightly acute (Fig. 5O). Trochanters with acute internal angle (Figs 5Q, 6C), metatibial spurs equal and short (Fig. 6F). Last sternite abruptly curved (Fig. 6P). Male genitalia with long and triangular parameres, phallus slightly widened at base and curved at apex (curvature of about 90°), with transparent keel on ventral part, phallus with thorns modified to receive parameres (Fig. 7K). Female genitalia with slender European Journal of Taxonomy 1022: 202–242 (2025) 224 stylus, coxites, and valvifers as independent structures, with slender coxites and S shape, valvifers not fused at base (Fig. 7N). Distribution Lyconotus is distributed in the Neartic and Neotropical regions. It has been reported in northeastern, southeastern and southern United States as well as in Mexico, Panama, and Colombia (Dugés 1878, 1896; Green 1949; Pérez-Hernández et al. 2019; GBIF 2023a). Genus Neolycus Bourgeois, 1883 Figs 4B, E–F, M–N, P, R–S, 5C, F, I-K, M, P, R–S, 6B, E, K, M, T, 7A–B, G, J, L, 9D–E Neolycus Bourgeois, 1883: 61. Type species Lycus schoenherri Chevrolat, 1834. Diagnosis Neolycus can be distinguished from Lycomesus, Lyconotus, Lycorectus gen. nov., and Rhyncheros by the presence of sexually dimorphic elytra, males with elytra considerably dilated in the middle, females with moderately dilated elytra (Fig. 9D–E); membranous wing with the radiomedial loop broadly curved (Fig. 5M); the metatibial spurs unequal, the inner acute, the outer slightly broader apically and bluntly rounded, and situated close to each other at the base (Fig. 6E); the aedeagus with thorns on the phallus (Figs 6T, 7A–B, G, J). In female genitalia, the valvifers are independent structures fused at the base (Fig. 7L). Material examined Refer to Supp. file 1. Redescription Body slender. Head mostly concealed by pronotum, rostrum variable in length (often very long) (Fig. 4B). Interantennal distance less than 0.09 mm. Antennomere III longer than antennomere IV (Fig. 4E–F). Variable labrum shape (semi-square to rectangular) (Fig. 4M). Mandibles as long as labrum. Pronotum widest at base, with variable anterior border shape (rounded to strongly angulated) (Fig. 4R–S), and anterior margins not prominent, pronotal margin longer than 0.82 mm. Elytra with sexual dimorphism, male with considerably dilated elytra, female with moderately dilated elytra (Fig. 9D–E), both sexes with four distinct costae on each elytron. Radiomedial loop broadly curved (Fig. 5M). Trochanters with rounded internal angle (Figs 5P, R–S, 6B), metatibial spurs unequal (inner acute, outer slightly broader apically and bluntly rounded) and long (Fig. 6E). Male with straight last sternite. Male genitalia with variable shape of parameres (moderately short, or very long and triangular) (Figs 6T, 7A), phallus slightly widened at base and straight to apex (Fig. 7B, G), with a small and transparent keel on ventral part; phallus with thorns of variable length (Figs 6T, 7A–B, G, J). Female genitalia with elongated stylus, coxites, and valvifers as independent structures, with coxites elongated and robust, valvifers fused at base (Fig. 7L). Distribution Neolycus is distributed sympatrically with Rhyncheros throughout the Neartic and Neotropical regions, with records from southern United States and Mexico (Dugés 1878, 1896; Gorham 1880, 1884; Green 1949; Pérez-Hernández et al. 2019; GBIF 2023b), and Peru (Bocak et al. 2015). GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 225 Genus Rhyncheros LeConte, 1881 Figs 4D, H–J, T–U, W, 5A–B, E, N, 6A, D, G–I, O, Q–R, 7C, F, H–I, M, 9F–G Rhyncheros LeConte, 1881: 17. Thoracocalon Bourgeois, 1883: lxi. Type species Lycus sanguinipennis Say, 1823. Diagnosis General appearance similar to Lycorectus gen. nov. but differs in the shape of the abdominal sternites with acute posterior angles; in males the posterior border of the seventh sternite is concave or emarginate (Fig. 6H–I); aedeagus with parameres of variable length, phallus widened at base, straight to slightly Fig. 9. Dorsal view of the North American Lycini Laporte, 1836. A. Lycomesus llorentei ZaragozaCaballero & González-Ramírez, 2019, ♂ (CNIN, QR 70969). B. Lyconotus semiustus (Chevrolat, 1834), ♂ (CNIN, QR 68732). C. Lyconotus semiustus (Chevrolat, 1834), ♀ (CNIN, QR 68736). D. Neolycus schoenherri (Chevrolat, 1834), ♂ (CNIN, QR 67812). E. Neolycus schoenherri, ♀ (CNIN, QR 67829). F. Rhyncheros loripes (Chevrolat, 1834), ♂ (CNIN, QR 68174). G. Rhyncheros loripes, ♀ (CNIN, QR 68175). European Journal of Taxonomy 1022: 202–242 (2025) 226 curved at apex, in some species, the phallus lacks thorns, in other instances, however, these structures are clearly present (Fig. 7C, F, H–I). Material examined Refer to Supp. file 1. Redescription Body slender. Head mostly concealed by pronotum, rostrum variable in length (moderately short, often very long) (Fig. 4H–J). Interantennal distance variable in length (less than 0.09 mm to more than 0.13 mm) (Fig. 4D). Antennomere III longer than antennomere IV. Variable labrum shape (semicircular, semi-square to rectangular) (Fig. 4L). Mandibles as long as labrum. Pronotum widest at base, with variable anterior border shape (rounded to strongly angulated) (Figs 4W, 5A–B), and variable anterior margins shape (not prominent to prominent) (Fig. 4T–U), wide pronotal margin (0.53–0.81 mm, often more than 0.82 mm). Elytra without sexual dimorphism, with four distinct costae on each elytron (Fig. 9F–G). Radiomedial loop slightly curved (Fig. 5N). Trochanters with rounded internal angle (Fig. 6A), metatibial spurs equal, acute and long (Fig. 6D, G). Male with straight last sternite. Male genitalia with variable shape of parameres (moderately long, often short), phallus slightly widened at base, straight to slightly curved at apex, with diminutive and transparent keel on ventral part, phallus thorns may or may not be present and variable in length (Fig. 7C, F, H–I). Female genitalia with variable length of stylus, coxites and valvifers fused, with robust coxites and elongated shape, valvifers not fused at base (Fig. 7M). Distribution Rhyncheros is distributed in the Neartic and Neotropical regions. It has been reported in southern United States as well as in Mexico (Dugés 1878, 1896; Gorham 1880, 1884; Green 1949; Pérez-Hernández et al. 2019; GBIF 2023b). Genus Lycorectus González-Ramírez & Zaragoza-Caballero gen. nov. urn:lsid:zoobank.org:act:82069CAA-EF3B-4811-915C-44A4EE7F76F0 Figs 10–11 Type species Lycostomus sordidus Gorham, 1880 (Figs 10–11), by present designation. Diagnosis Lycorectus gen. nov. is closely related to Rhyncheros. In both genera, the prosternum is strongly carinated, the mesosternum is slightly prominent, the internal angles of the metacoxae are blunt, and the metatibial spurs are equal and long. The main differences between Rhyncheros and Lycorectus are in the shape of the posterior border of the seventh sternite in males, which may be either concave or emarginate in Rhyncheros, but is slightly convex in Lycorectus. In addition, the phallus of Rhyncheros may or may not have thorns, whereas the phallus of Lycorectus lacks thorns (Fig. 10C–E). Etymology The name Lycorectus is derived from the Greek term ‘Lykos’ = ‘wolf’ and Latin term ‘rectus’ = ‘straight’. The latter pertains to the straight shape of phallus of the aedeagus in comparison to the same structure in Lycomesus and Lyconotus. GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 233 Our results contrast with those of Kusy et al. (2020), who examined the genitalia of the type species of Rhyncheros (R. sanguinipennis) and Lyconotus (Lycus lateralis) and suggested them to be closely related. Therefore, after comparing the type species and analysing the relationships between the Nearctic species, they concluded that Lyconotus should be regarded as a synonym of Rhyncheros (Kusy et al. 2020). However, male genital characters have been utilized to distinguish Lyconotus from other genera of Lycini (Green 1949; Zaragoza-Caballero 1995; Zaragoza-Caballero & González-Ramírez 2019). Examples of such characters include: parameres about one-third length of the phallus and phallus sharply bent downwards medially (Fig. 7K) (Green 1949; Zaragoza-Caballero 1995; Zaragoza-Caballero & González-Ramírez 2019). Our analysis identified the last character as a synapomorphy supporting the monophyly of Lyconotus (Fig. 8). Furthermore, Green (1949) and Zaragoza-Caballero (1995) suggested that additional characteristics, such as the presence of spinose trochanters (Figs 5Q, 6C) and the male last sternite abruptly bent medially (Fig. 6P), can be utilized to distinguish these genera from other North American Lycini. In our analysis, we identified the latter as a synapomorphy that supports the Lyconotus clade (Fig. 8). Our phylogenetic analysis corroborates the previous morphological considerations, so we consider Lyconotus as a valid genus. The two analysed species of Lycostomus were recovered in distinct clades (Fig. 8): L. praeustus was recovered as sister to (Lycostomus rubrocinctus) + (Lycus + (Lycorectus + (Neolycus + Rhyncheros). Therefore, it would be possible to transfer L. praeustus to another genus; however, before making any taxonomic change in this regard, it would be prudent to broaden the sampling of Asian Lycostomus. We prefer to wait because Kusy et al. (2020) found that the Asian Lycostomus represent a monophyletic group. In our analysis, the genus Lycus is sister to the rest of the North American Lycini (Fig. 8). Lycus was recovered as monophyletic, with low statistical support; however, it was supported by one synapomorphy (Table 3; Supp. file 4: fig. S2): bifurcated phallus apex (character 79: state 2; Fig. 7D), and a combination of the following homoplastic characters (Supp. file 4: fig. S2): male with bidentate antennomeres IV–X (2:0), prementum with median suture (21:1), angulated anterior margin of pronotum (22:1); punctuated surface on pronotal lateral margin (26:0), sparse pronotal pubescence on lateral margins (27:0), transverse sternellum with posterior rami (39:0), apex of elytra rounded (44:1; Fig. 5G). The support (UFBoot = 91; PP = 0.83) for the monophyly of Lycus is consistent with the molecular phylogeny of Kusy et al. (2020); however, as with Lycostomus, it is necessary to consider a larger sample to analyse the relationships in the genus with greater rigour. Our cladogram recovered a clade that comprises the remaining North American Lycini (Lycorectus, Neolycus and Rhyncheros). This clade was supported by three synapomorphies (Table 3; Supp. file 4: fig. S2): strongly carinated prosternum (character 36: state 1), strongly protuberant mesosternum (50:2), and fused coxites-valvifers (83:2). Nonetheless, the diagnostic usefulness of theses synapomorphies is weakened by several reversals within the clade (Supp. file 4: fig. S2). The clade was additionally supported by the following combination of homoplastic character states (Supp. file 4: fig. S2): labrum with lateral margins semi-rounded (character 12: state 1), mandible as long as labrum (14:1; Fig. 4L–M), pronotum with wide longitudinal areolar, spindle-shaped (32:2), prosternum not protuberant (34:0), prosternum with rounded posterior border (38:0), female with last sternite cleft (66:2), and coxites longer than valvifers (88:2). The character state of mandibles as long as labrum (14:1; Fig. 4L–M) has an independent origin (with respect to Lycostomus praeustus) (Supp. file 4: fig. S2); however, its occurrence within this clade is due to common ancestry. Lycorectus sordidus gen. et comb. nov. was recovered as sister to Neolycus and the other species in the genus Rhyncheros. The species L. sordidus can be characterized by one apomorphic character (Table 3; Supp. file 4: fig. S2): male six sternite with convex posterior margin (character 63 state 5; Fig. 6L), European Journal of Taxonomy 1022: 202–242 (2025) 234 and additionally 14 homoplastic characters (Supp. file 4: fig. S2): male with rectangular antennomeres (character 2: state 2), length of fourth maxillary palpomere shorter than second maxillary palpomere (17:0), male pronotum with short anterior angles (23:1), lateral margins of pronotum scarcely pubescent (27:0), elytra dilated towards apex (40:2; Figs 11A), female elytra dilated width-humeral width ratio 1.7 to 2.1 (42:1), bipartite scutellum apex (48:2), narrow mesosternum (51:2), metacoxae inner angle acute in both male and female (53:2; 54:2), in male genitalia, apex of parameres acute (71:1; Fig. 11E), phallus shape as in morphological group 1 (72:1; Figs 3, 11C–E), length of the stylus is greater than 0.30 mm (80:2), and coxites with medial emargination (85:1). Our study is the first to use L. sordidus in a phylogenetic context. We proposed the description of a new genus for L. sordidus based on the IW topology, which shows it is outside the North American Rhyncheros clade; the apomorphic character recovered in our analysis (male convex posterior edge on seventh sternite) (63:5; Fig. 6L) and on the fact that it has been reported for North and South America (Gorham 1880; Pérez-Hernández et al. 2019). In the results section, we provide a description for this new genus. Nonetheless, it is imperative to assess the phylogenetic relationships of the South American species of Lycini because L. sordidus shares diagnostic characters with Thoracocalon (pronotum distinctly rounded in the males and broadly foliated lateral margins), a subgenus that previously included only South American Lycini (Gorham 1880; Bourgeois 1883, 1889, 1901; Pic 1922). Based on our results, Neolycus and Rhyncheros are sister taxa (Fig. 8). This group was supported by the combination of the following homoplastic characters (Supp. file 4: fig. S2): labrum wider than longer (character 10: state 1), pronotum with rounded lateral margins (25:1), phallus with pair of thorns (73:1; Figs 6T, 7A–C, F–J), and length of the stylus between 0.22 mm to 0.29 mm (80:1). With respect to thorns on the phallus (character 73: state 1), Kusy et al. (2020) considered the absence of thorns in middle part of phallus as a diagnostic character of Rhyncheros. We do not support that opinion, because our results showed that this condition was lost in only two species of Rhyncheros (R. nigrofumosus and R. loripes), and hence it is homoplastic (Supp. file 4: fig. S2). Neolycus is the second most diverse genus of the American Lycini (seven species) and is distributed across both the Nearctic and Neotropical regions. However, its greatest diversity has been recorded in the Neotropical region (Dugés 1878, 1896; Gorham 1880, 1884; Green 1949; Bocak et al. 2015; PérezHernández et al. 2019; GBIF 2023b). In our IW cladogram, Neolycus was recovered as monophyletic although with low statistical support (BS = 59; JS = 64; UFBoot = 80) (Fig. 8). Despite the low support, this result is consistent with the molecular phylogenetic analysis performed by Kusy et al. (2020). In our topology, Neolycus was supported by one apomorphy (Table 3; Supp. file 4: fig. S2): dissimilar metatibial spurs (the inner spur has an acute apex, while outer spur is slightly broader at apex and bluntly rounded) (character 61: state1; Fig. 6E), and by a combination of the following homoplastic character states (Supp. file 4: fig. S2): prementum with a median suture (21:1), posterior margin of last sternite emarginate (66:1), female genitalia, with coxites and valvifers not joined (83:0); and valvifers fused at base (89:1) (Fig. 7K). We confirm Green’s (1949) use of dissimilar metatibial spurs as a diagnostic character for Neolycus. The present study is the first to incorporate characters related to female genitalia, although these characters are homoplastic, they add support for the monophyly of Neolycus and contribute to the overall resolution of the tree. According to our topology, Rhyncheros is supported as monophyletic, and can be circumscribed by one synapomorphy: labrum semi-circular (character 9: state 1), and by the combination of the following homoplastic characters states (Table 3; Supp. file 4: fig. S2): labrum with rounded lateral margins (12:2), securiform last maxillary palpomere (15:2), fourth maxillary palpomere dilated at the apex (16:1), securiform last labial palpomere (18:2), and spatula-shaped scutellum (46:1). The maxillary palpi character state (15:2) has two independent origins (with respect to Calopteron Laporte, 1838) (Supp. file 4: fig. S2). However, its presence within Lycostomus rubrocinctus and Rhyncheros is a result GONZÁLEZ-RAMÍREZ M. et al., Systematic of North American Lycini (Coleoptera: Lycidae) 235 of common ancestry. The hypothesis that Calopteron is the sister group of Lycini has been proposed by various works (Bocak & Bocakova 1990; Bocakova 2003, 2005; Bocak & Bocakova 2008). This hypothesis is coherent because the larvae of Calopteron, Lycostomus and Rhyncheros share numerous morphological traits, for example, it has been reported that the larvae of these genera have maxillary palpi that are three-segmented and longer than mala (McCabe & Johnson 1979; Bocak & Matsuda 2003; González-Ramírez & Zaragoza-Caballero 2024). The genus Rhyncheros is the most diverse among the American lycines, consisting of 29 species, and is widely distributed in the Nearctic and Neotropical regions (Supp. file 5: fig. S1) (Dugés 1878, 1896; Gorham 1880, 1884; Kleine 1933; Blackwelder 1945; Green 1949; Pérez-Hernández et al. 2019; GBIF 2023b). According to Kusy et al. (2020) the genus was corroborated as monophyletic and is the sister group to Neolycus, and our results support this. Our analysis recovered to Rhyncheros godmani as a sister group to other members of the genus Rhyncheros. Furthermore, the analysis yielded the identification of two subclades within Rhyncheros (Fig. 8). The first is formed by R. nigrofumosus, R. fuliginosus and R. carnifex, which can be characterized by the unique combination of four homoplastic characters (Supp. file 4: fig. S2): labrum with anterior margin emarginate (character 11: state 2), lateral margin of pronotum abundant pubescence (27:2), sternellum V-shaped with short branches (39:2), and males with metacoxae inner angle acute (53:2). The second subclade includes R. sanguinipennis (type species) and most of the species in this genus (9): R. sanguineus, R. lineicollis, R. fulvellus, R. rusticus, R. fulvellus femoratus, R. simulans, R. sagittatus, R. loripes, and R. minutus. This group of species can be characterized by the unique combination of four homoplastic characters (Supp. file 4: fig. S2): rectangular labrum in dorsal view (character 9: state 3), prosternum with posterior border truncated (38:1), male elytra dilatated width-humeral width radio less than 1.79 (41:0), and coxites with distant inner apical margin (87:1; Fig. 7M). Rhyncheros occurs sympatrically with Neolycus in the Nearctic and Neotropical regions (Gorham 1880; Green 1949; Pérez-Hernández et al. 2019; Kusy et al. 2020). Our research represents a significant step in the understanding of the evolution of North American lycines, because it incorporates a representative sample of taxa, the largest so far, and incorporates an extensive revision of morphological characters in a phylogenetic context. According to our morphological phylogenetic hypothesis, Lycini are supported as monophyletic. Our results are in a better agreement with a natural classification considering five genera for the North American Lycini, which requires the reinstatement of Lyconotus, and the newly proposed genus Lycorectus. Ours results extend the classification of North American lycines presented by Kusy et al. (2020) to include Lycomesus, Lyconotus, Lycorectus gen. nov., Neolycus and Rhyncheros. The phylogeny of North American Lycini is derived exclusively from specimens found in a museum collection, thus highlighting the crucial role of scientific collections in evolutionary studies. Based on the study of this material, the examination of male and female genitalia has yielded a comprehensive understanding of the morphology of North American lycines and offers significant taxonomic value for diagnosing genera. Additionally, the characters obtained from both male and female genitalia provide support and contribute to the resolution of clades in our analysis. Future research on Lycini should prioritize the utilization of molecular and morphological evidence from North and South American lycines to examine the impacts of alternative data and establish a more robust phylogenetic hypothesis for the South American taxa within Lycini. Acknowledgments This paper is part of the requirements for obtaining a Doctoral degree at the Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México (UNAM). We thank the Consejo Nacional de Humanidades, Ciencia y Tecnología (CONAHCyT) for the support of this research through funding European Journal of Taxonomy 1022: 202–242 (2025) 236 and with a graduate scholarship to the first author. We appreciate the help of the curator Michael Geisel for granting us access to the Coleoptera collection at the Natural History Museum (NHMUK). Additionally, we thank Keita Matsumoto and Miriam Aquino Romero for capturing photographs of the type species of Lycini at the NHMUK, to Jon Gelhaus for allowing us access to material of Rhyncheros sanguinipennis at the Academy of Natural Sciences in Philadelphia (ANSP), and to Jason D. Weintraub for photographing the specimen; to Chris Grinter and Rachel Diaz-Bastin for providing access to material and photographs of the species Rhyncheros nigrofumosus and Lyconotus lateralis. We thank Geovanni M. Rodríguez Mirón, Viridiana Vega Badillo, and Francisco Armendáriz Toledano for helping to improve the manuscript. 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Dugesiana 26 (2): 99–102. https://doi.org/10.32870/dugesiana.v26i2.7075 Printed versions of all papers are deposited in the libraries of three of the institutes that are members of the EJT consortium: Muséum national dʼHistoire naturelle, Paris, France; Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural Sciences, Brussels, Belgium. The other members of the consortium are: Meise Botanic Garden, Meise, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain; Leibniz Institute for the Analysis of Biodiversity Change, Bonn – Hamburg, Germany; National Museum of the Czech Republic, Prague, Czech Republic; The Steinhardt Museum of Natural History, Tel Aviv, Israël. Supplementary files Supp. file 1. List of specimens examined. https://doi.org/10.5852/ejt.2025.1022.3089.13769 Supp. file 2. Data matrix for phylogenetic analysis of the tribe Lycini Laporte, 1836 (Lycidae: Lycinae). https://doi.org/10.5852/ejt.2025.1022.3089.13771 Supp. file 3. Strict consensus tree of parsimony analysis under equal weight. https://doi.org/10.5852/ejt.2025.1022.3089.13773 Supp. file 4. A single cladogram of parsimony analysis under implied weight. Fig. S1. Unambiguous character changes mapped on branches in WinClada. Fig. S2. Slow optimization using delayed (DELTRAN) transformation. Fig. S3. Fast optimization using accelerated (ACCTRAN). https://doi.org/10.5852/ejt.2025.1022.3089.13775 Supp. file 5. Phylogenetic tree of North American Lycini Laporte, 1836, based on the maximum likelihood analysis. https://doi.org/10.5852/ejt.2025.1022.3089.13777 Supp. file 6. Phylogenetic tree of North American Lycini Laporte, 1836, based on the Bayesian inference. https://doi.org/10.5852/ejt.2025.1022.3089.13781