Introduction
Telegeusinae are the most diverse subfamily of Omethidae (Coleoptera), a heterogeneous and soft-bodied family within Elateroidea (Kundrata et al. Reference Kundrata, Bocakova and Bocak2014; Cai et al. Reference Cai, Tihelka, Giacomelli, Lawrence, Ślipiński and Kundrata2022). The subfamily is found only in the Western Hemisphere, from the southern United States of America to northern South America (Miller Reference Miller, Arnett, Thomas, Skelley and Frank2002; Roza et al. Reference Roza, Constantin and Mermudes2019; Ferreira Reference Ferreira2025; Fiuza et al. Reference Fiuza, Roza and Biffi2025; López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2026)). Telegeusinae include six genera and 28 described species, with the majority of the group’s diversity occurring in Mexico (Pérez-Flores and Zaragoza-Caballero Reference Pérez-Flores and Zaragoza-Caballero2024; Ferreira Reference Ferreira2025). The group has received some attention in recent years, with several new species and new genera described (e.g., Roza et al. Reference Roza, Constantin and Mermudes2019; López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021; Ferreira Reference Ferreira2025). Despite this effort, the group is still largely unknown, with no larva or female (which are hypothesised to be paedomorphic) described and with several countries within the distributional range of Omethidae lacking records of the family.
I received a loan of unidentified phengodid beetles from the Canadian Museum of Nature, Ottawa, Ontario, Canada (CMN), and the zoological collection of the Museum of Biology of Lund, Lund, Sweden (MZLU), which contained several Telegeusinae specimens. Among them, two specimens were noticeably distinct from the more common Telegeusis Horn, 1895. Based on the most updated keys for genera (López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021; Ferreira Reference Ferreira2025), the specimens were identified as Stenodrilus López-Pérez and Zaragoza-Caballero, Reference López-Pérez and Zaragoza-Caballero2021, a monotypic genus from southern Mexico, but they differed significantly from the only described species. In this paper, I describe these two specimens as two new species. These species represent the first Omethidae known from Honduras and the first record of Stenodrilus for Central America. I also provide an identification key and a distributional map for Stenodrilus species, as well as the first description of the terminalia for the genus. Finally, I discuss the morphology of the genus and new interpretations of some of its diagnostic characters, and I provide a new diagnosis for Stenodrilus.
Material and methods
The specimens examined in this study are deposited at the CMN and MZLU. They were identified based on the most recent generic keys for the family (López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021; Ferreira Reference Ferreira2025). Labels are cited in the following fashion: lines on the same label are separated by a slash “/”, and separate labels are indicated by a double backslash “\\”. Additional information provided by me appears between square brackets “[]”. Words printed on the labels in uppercase are transcribed as such. Specimen count appears between parentheses after label information. Collection acronyms appear after all labels for the specimens. Country (bold, uppercase) and province or state (italic, lowercase) information appears before the label information.
For the classification, I follow Cai et al. (Reference Cai, Tihelka, Giacomelli, Lawrence, Ślipiński and Kundrata2022). The general terminology follows López-Pérez and Zaragoza–Caballero (Reference López-Pérez and Zaragoza-Caballero2021). For the head morphology, I follow Miller (Reference Miller, Arnett, Thomas, Skelley and Frank2002), calling the free lobe between the mandibles the labrum (instead of clypeus, as in López-Pérez and Zaragoza-Caballero (Reference López-Pérez and Zaragoza-Caballero2021)). For the hind wings, I follow Lawrence et al. (Reference Lawrence, Zhou, Lemann, Sinclair and Ślipiński2021). I detached the wing and abdomen from both specimens, and I dissected the latter. Before dissection, the abdomen was placed in a 10% KOH solution for about 12 hours at room temperature. Afterwards, the abdomen was rinsed in distilled water, dissected, and photographed. The dissected abdomen, terminalia, and genitalia are permanently preserved in glycerine in a microtube, and the wing is glued on a card, with both preparations placed on the pin under each respective specimen.
Photographs and measurements were taken with a Leica DFC450 and Application Suite CV3 multifocus software (Leica Microsystems, Wetzlar, Germany). Total length was measured as the sum of the head, thorax, and abdominal (up to hind wing apex) lengths in lateral view because extracting only overall body size could be misleading due to pronotal and head declination. The photographs were edited using Adobe Photoshop, and the plates were designed with Adobe Illustrator (Adobe Systems, Adobe, San Jose, California, United States of America). The map was generated using the software Quantum GIS, version 2.14.3 (QGIS Development Team 2023).
Results
Stenodrilus López-Pérez and Zaragoza-Caballero, 2021
Stenodrilus howdenorum new species holotype: A, habitus, dorsal; B, habitus, lateral; C, head, dorsal; D, pronotum, dorsal; E, head, frontal, with drawing of the labrum shape; F, head, lateral; and G, hind wing. Scale bar: A–B, 1 mm; C–F: 0.5 mm; G: 1 mm.
Stenodrilus howdenorum new species holotype: A, abdomen, dorsal; B, abdomen, ventral; C, tergite IX and X; D, ventrite 9; E, aedeagus, dorsal; F, aedeagus, lateral; and G, aedeagus, ventral. Scale bar: A–B, 0.5 mm; C–G: 0.2 mm.
Stenodrilus megalocephalus new species holotype: A, habitus, dorsal; B, habitus, lateral; C, head and pronotum, dorsal; D, head, frontal, with drawing of the labrum shape; E, head, lateral; F, head, ventral; and G, hind wing. Scale bar: A–B, 1 mm; C–F: 0.5 mm; G: 1 mm.
Stenodrilus megalocephalus new species holotype: A, abdomen, dorsal; B, abdomen, ventral; C, tergite IX and X; D, ventrite 9; E, aedeagus, dorsal; F, aedeagus, lateral; and G, aedeagus, ventral. Scale bar: A–B, 0.5 mm; C–G: 0.2 mm.
Distribution of Stenodrilus species.
Schematic drawing of Telegeusinae aedeagus, with the entire piece and also each structure isolated, using Stenodrilus megalocephalus new species as a model. Scale bar: 0.2 mm.
Type species. Stenodrilus rodolfoi López-Pérez and Zaragoza-Caballero, Reference López-Pérez and Zaragoza-Caballero2021
Diagnosis. Head with postocular area coarsely and deeply punctate laterally (Figs. 1F, 3E). Antennae filiform with 11 antennomeres (Figs. 1A, 3A). Labrum with medium tooth of variable length. Maxillary palpi with three palpomeres, last palpomere enlarged and flattened, 2–2.5 times as long as the two preceding palpomeres’ combined length, about three times longer than labial palpi’s combined length. Labial palp with two palpomeres, last palpomere digitiform, as long as preceding palpomere. Tentorium with two contiguous pits; gula with two obliquely separate sutures (Fig. 3F). Mesotibial spurs smooth. Wing with radius anterior vein (RA) well marked, radial cell (R) absent, radial cross-vein 3 long, oblique in relation to RA, radial cross-vein 4 absent, radius posterior vein (RP) present, close to half of media posterior vein 1 + 2 (MP1 + 2) length, medial field venation reduced, with only media posterior vein 3 + 4 (MP3 + 4) and the cubitoanal strut (CAS) distinct, anal posterior vein 3 + 4 (AP3 + 4) distinct, jugal vein (J) absent (Figs. 1G, 3G). Abdominal tergites and ventrites well sclerotised (Figs. 2A–B, 4A–B); tergite IX without any division, around two times as long as tergite X, which is slightly convex dorsally; sternite IX elliptical, well sclerotised (Figs. 2C–D, 4C–D). Aedeagus with phallobase large, covering nearly two-thirds of aedeagus length ventrally; median lobe cylindrical, curved at base and near the apex, which is rounded and not constricted apically; parameres parallel, twisted near apex, which is rounded, spatulate, and granulate, with long semierect setae, and with a subbasal medial arch that encircles the median lobe posteriorly and extends almost to the parameres’ apex (Figs. 2E–G, 4E–G, 6).
Comparison to other Telegeusinae genera. This genus can be distinguished from Ceratogeusis Ferreira, 2025 by the filiform antennae (flabellate antennae from antennomere 5–10 in Ceratogeusis), the labial palpi with two palpomeres (one palpomere in Ceratogeusis), and tentorium with two contiguous pits (a single pit in Ceratogeusis); from Platyphalus López-Pérez and Zaragoza-Caballero, 2026 by the head postocular area coarsely and deeply punctate laterally (postocular area feebly punctate laterally in Platyphalus), tentorium with two contiguous pits (a single pit in Platyphalus), and gula with two sutures (one suture in Platyphalus); from Pseudotelegeusis Wittmer, 1976 by the filiform antennae (serrate in Pseudotelgeusis) and the labial palpi with two palpomeres (one palpomere in Pseudotelegeusis); from Pseudokarumia Pic, 1931 by the smooth mesotibial spurs (serrate mesotibial spurs in Pseudokarumia); and from Telegeusis Horn, 1895 by the apical labial palpomere digitiform, as long as the preceding palpomere (apical labial palpomere enlarged and flattened, much longer than preceding palpomeres in Telegeusis).
Remarks. Some characters previously assigned as diagnostic for Stenodrilus do not seem to be good distinguishable features when compared to the rest of the subfamily members. The genus was originally described as having a single tentorial pit (López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021), but upon closer examination of the two new species and of illustrations of S. rodolfoi, it is clear that the species in the genus actually have two contiguous tentorial pits, barely distinguishable from each other (Fig. 3F; López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021: Fig. 2D). To date, the only known genera with a single tentorial pit are Ceratogeusis and Platyphalus. Although Pseudotelegeusis jiliotupaensis Zaragoza-Caballero, 2008 is also described with a single tentorial pit, that species is most likely assigned to the wrong genus (Roza et al. Reference Roza, Constantin and Mermudes2019).
A character raised by López-Pérez and Zaragoza-Caballero (Reference López-Pérez and Zaragoza-Caballero2021, Table 1) as diagnostic for Stenodrilus and Platyphalus is the maxillary palpi with only three palpomeres (in contrast to four palpomeres in all other Telegeusinae genera). This character, however, is most likely erroneously described for all the other genera in the subfamily. Telegeusinae maxilla has a palpifer of variable distinction, which can be mistakenly identified as a first palpomere. In addition, the last maxillary palpomere, which is always enlarged and flattened, has a proximal region shaped like the other palpomere (more-or-less cylindrical) and can be erroneously assumed to be a preapical palpomere.
In this context, several species of Telegeusis are already described with only three maxillary palpomeres (Pérez-Flores and Zaragoza-Caballero Reference Pérez-Flores and Zaragoza-Caballero2024) instead of the four usually assigned to this structure in the genus. The maxillary palpi illustrated in Roza et al. (Reference Roza, Constantin and Mermudes2019, Fig. 3A) for Pseudotelegeusis meloi is clearly composed of three palpomeres, although mistakenly described with four palpomeres (the first “palpomere” is actually the palpifer in dorsal view). In addition, all species of Telegeusis and Pseudotelegeusis examined during the present study have only three maxillary palpomeres. A complete morphological revision of the group is needed to resolve these inconsistencies and diagnose each of the known genera correctly.
Another character raised for Stenodrilus in the original description (López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021) is the somewhat slender parameres in the genitalia. The aedeagus of the genus seems to be consistent with what is found in other genera in the subfamily. The shape, length, and width of the parameres probably present species-level variations rather than generic diagnostic characters (see more details in the Discussion section).
Stenodrilus howdenorum Roza new species
ZooBank Nomenclature Act #: urn:lsid:zoobank.org:act:81FCCD12-394E-4F92-BD47-03F2531BEE42
Type material. Holotype. HONDURAS: Francisco Morazán. “HONDURAS: F. Morazán/30kmE Tegucigalpa/Cerro Uyuca/3.VI.1994 1800m/H and A Howden\\ Malaise Trap” (1 male; CMN).
Diagnosis. Body overall dark brown, with lighter legs (Fig. 1A, B). Head slightly longer than wide. Eyes small, occupying about one-sixth of head width in dorsal view (Fig. 1C); in lateral view, posterior margin vertical, straight; postocular length twice the eye length in lateral view (Fig. 1F). Labrum without prominent central teeth (Fig. 1E). Pronotum slightly wider than long, anterior margin straight, posterior margin obliquely projected medially (Fig. 1D). Elytron around 4.5 times as long as wide (Fig. 1A).
Description. Measurements. Total length: 4.78 mm. Head length: 0.77 mm. Head width: 0.64 mm. Pronotum length: 0.50 mm. Pronotum maximum width: 0.57 mm. Elytron length: 1.67 mm. Elytron maximum width: 0.37 mm.
Morphology. Head slightly longer than wide, 1.2 times as long as wide, integument glossy, smooth dorsally and ventrally, shagreened laterally, coarsely setigerous punctate, with shallow punctures dorsally and ventrally and deep punctures laterally, eyes small, slightly protruding, occupying around one-sixth of head width in dorsal view (Fig. 1C), postocular length twice the eye length in lateral view, eye posterior margin straight (Fig. 1F); labrum without prominent central teeth (Fig. 1E); maxilla with last palpomere almost three times as long as the two preceding palpomeres combined; labium with last palpomere as long as the preceding palpomere; tentorium with two contiguous pits; gula with two obliquely separate sutures. Pronotum slightly wider than long, integument glossy and smooth, heavily setigerous punctate with shallow and almost indistinct punctures, anterior margin straight, lateral margins dorsally subparallel, posterior margin obliquely projected medially (Fig. 1D). Elytron heavily setigerous punctate, around 4.5 times as long as wide (Fig. 1A). Posterior wings with venation as in the genus description (Fig. 1G). Abdomen with tergite and ventrites transverse (Fig. 2A, B). Tergite IX transverse, 1.2 times as wide as long in the middle line; tergite X 1.4 times as long as wide (Fig. 2C); ventrite 8 with posterior margin straight (Fig. 2B); ventrite 9 1.4 times as long as wide, posterior margin rounded (Fig. 2D). Aedeagus with phallobase large, elongate, covering nearly two-thirds of aedeagus length ventrally; median lobe cylindrical, slightly curved at base and almost straight near apex, which is rounded and not constricted apically; paramere parallel, about four times as long as wide, twisted near rounded, granulate apex, with long semierect setae (Fig. 2E–G).
Colouration. Body overall dark brown, with lighter legs (Fig. 1A, B).
Etymology. The name recognises two accomplished entomologists, Henry Howden (1925–2014) and Anne Howden (1927–2016), who contributed tremendously to the knowledge of world beetles. They gathered an enormous collection that was later donated to the Canadian Museum of Nature. Their collection continues to house several undescribed taxa, particularly from the Neotropics. The name is Latin in the genitive, plural.
Natural history. The holotype was collected at the beginning of June, at the beginning of the rainy season in Honduras. Specimens of this species have not been seen alive, and no information regarding its habits is known. Cerro Uyuca, the type locality, is primarily composed of pine–oak forests.
Distribution. Stenodrilus howdenorum is known to occur only at mid-elevation (1800 m) of Cerro Uyuca, Francisco Morazán Department, Honduras (Fig. 5).
Remarks. Stenodrilus howdenorum can be easily differentiated from Stenodrilus megalocephalus new species by the short head, only slightly longer than wide (distinctively long head, about 1.4 times as long as wide in Stenodrilus megalocephalus), the larger eyes, covering around one-sixth of the head width in dorsal view (small, feebly protruding eyes, occupying around one-tenth of the head width in dorsal view in Stenodrilus megalocephalus), the postocular length twice the eye length in lateral view (2.5 times the eye length in lateral view in Stenodrilus megalocephalus), and the median lobe apex almost straight (distinctly curved in Stenodrilus megalocephalus). Stenodrilus howdenorum can be differentiated from S. rodolfoi by the dark brown head and pronotum (yellowish brown in S. rodolfoi) and by the elytron 4.5 times as long as wide (4 times as long as wide in S. rodolfoi).
Stenodrilus megalocephalus Roza new species
ZooBank Nomenclature Act #: urn:lsid:zoobank.org:act:5BA0CC35-88F7-44ED-8F82-E23C3877A2BE
Type material. Holotype. HONDURAS: Yoro. “Honduras: Yoro,/Palo de Comba/15°11'N, 87°39'W/31.VIII.1995 leg. R. Cave\\ Malaise trap in/mid elevation/secondary forest” (1 male; MZLU).
Diagnosis. Body overall dark brown, with paler legs (Fig. 3A, B). Head distinctly longer than wide. Eyes distinctively small, occupying about one-tenth of head width in dorsal view (Fig. 3C); in lateral view, posterior margin vertical, straight; postocular length 2.5 times the eye length in lateral view (Fig. 3E). Labrum with prominent central teeth (Fig. 3D). Pronotum slightly wider than long, anterior margin straight, posterior margin obliquely projected medially (Fig. 3C). Elytron around four times as long as wide (Fig. 3A).
Description. Measurements. Total length: 4.74 mm. Head length: 1.05 mm. Head width: 0.76 mm. Pronotum length: 0.69 mm. Pronotum maximum width: 0.72 mm. Elytron length: 1.63 mm. Elytron maximum width: 0.40 mm.
Morphology. Head distinctly longer than wide, 1.4 times the width, integument glossy, smooth dorsally and ventrally, shagreened laterally, coarsely setigerous punctate, with shallow punctures dorsally and ventrally, and with deep punctures laterally, eyes small, feebly protruding, occupying around one-tenth of the head width in dorsal view (Fig. 3C), postocular length 2.5 times the eye length in lateral view, eye posterior margin straight (Fig. 3E); labrum with prominent central teeth (Fig. 3D); maxilla with last palpomere about 2.5 times as long as the two preceding palpomeres combined; labium with last palpomere slightly longer than the preceding palpomere; tentorium with two contiguous pits; gula with two obliquely separate sutures (Fig. 3F). Pronotum slightly wider than long, integument glossy, smooth, heavily setigerous punctate with shallow and almost indistinct punctures, anterior margin straight, lateral margins dorsally subparallel, slightly divergent anteriorly, posterior margin obliquely projected medially (Fig. 3C). Elytron heavily setigerous punctate, around four times as long as wide (Fig. 3A). Posterior wings with venation as in the genus description (Fig. 3G). Abdomen with tergite and ventrites transverse (Fig. 4A, B). Tergite IX transversal, 1.3 times as wide as long in the middle line; tergite X 1.5 times as long as wide (Fig. 4C); ventrite 8 with posterior margin straight (Fig. 4B); ventrite 9 1.4 times as long as wide, posterior margin rounded (Fig. 4D). Aedeagus with phallobase large, elongate, covering nearly two-thirds of aedeagus length ventrally; median lobe cylindrical, strongly curved at base and near the apex, which is rounded and not constricted apically; parameres parallel, each about four times as long as wide, twisted near rounded, granulate apex, with long semierect setae (Fig. 4E–G).
Colouration. Body overall dark brown, with paler legs (Fig. 4A, B).
Etymology. The specific epithet is the junction of the Greek words mega (very large) and cephalus (head) and refers to the elongate head. The name should be treated as a noun in apposition.
Natural history. The holotype was collected at the end of August, close to the middle of the rainy season in Honduras. Specimens of this species have not been seen alive, and no information regarding its habits is known.
Distribution. Stenodrilus megalocephalus is known to occur only in mid-elevation secondary forest of Palo de Combas Village, Victoria Municipality, Yoro Department, Honduras (Fig. 5).
Remarks. Stenodrilus megalocephalus can be easily differentiated from the other two species in the genus by its distinctively long head, about 1.4 times as long as wide (versus head slightly longer than wide in Stenodrilus howdenorum and S. rodolfoi), the small, feebly protruding eyes, occupying around one-tenth of the width in dorsal view (occupying around one-sixth of the width in dorsal view in Stenodrilus howdenorum and S. rodolfoi), the postocular length 2.5 times the eye length in lateral view (twice the eye length in lateral view in Stenodrilus howdenorum and S. rodolfoi – although equivocally described as 2.5 times in S. rodolfoi (López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021)), and the median lobe apex distinctly curved (almost straight in Stenodrilus howdenorum and S. rodolfoi). It can be further differentiated from Stenodrilus howdenorum by the elytron four times as long as wide (4.5 times as long as wide in Stenodrilus howdenorum), and median lobe base distinctly curved (slightly curved in Stenodrilus howdenorum), and from S. rodolfoi by the dark brown head and pronotum (yellowish brown in S. rodolfoi).
Key to species of Stenodrilus López-Pérez and Zaragoza-Caballero, 2021
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1. Head distinctively long, about 1.4 times as long as wide; eyes small, feebly protruding, occupying around one-tenth of the head width in dorsal view; postocular length 2.5 times eye length in lateral view; aedeagus with median lobe apex distinctly curved…………………………………………… Stenodrilus megalocephalus new species
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–. Head short, slightly longer than wide; eyes large, covering around one-sixth of head width in dorsal view; postocular length twice the eye length in lateral view; aedeagus with median lobe apex almost straight…………………………………………… 2
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2. Head and pronotum dark brown; elytron 4.5 times as long as wide…………………………………………………… Stenodrilus howdenorum new species
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–. Head and pronotum yellowish brown; elytron four times as long as wide……………………………… Stenodrilus rodolfoi López-Pérez and Zaragoza-Caballero, Reference López-Pérez and Zaragoza-Caballero2021
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Discussion
Stenodrilus howdenorum and S. megalocephalus are the first telegeusine species recorded in Honduras, thereby improving current knowledge of the family in the taxonomically poorly known country. Many other species of the genus, from other locations and different countries in Central America, likely are scattered in entomology collections and awaiting description. The same goes for more populations of the currently described species. This paper sheds light on the distribution and richness of the subfamily and Stenodrilus, and it encourages other entomologists to find and describe more species of this rare group.
The new species described herein are mostly in agreement with the morphology of the type species, rendering the genus a general morphological homogeneity. When including the species described herein, the genus still has a clear diagnosis. It is interesting that the subfamily seems to have some extremely conserved morphological characters. The wing morphology, for instance, is exactly the same in all Stenodrilus species (López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021) and is also consistent with Pseudotelegeusis howdeni Wittmer, Reference Wittmer1976 (Wittmer Reference Wittmer1976) and several different Telegeusis species (Pérez-Flores and Zaragoza-Caballero Reference Pérez-Flores and Zaragoza-Caballero2024). Platyphalus is the only genus that presents some extra veins (López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021), and Pseudokarumia wing venation is not yet described.
On the other hand, the aedeagus is practically identical across the subfamily. With the exception of Pseudokarumia, all of the genera have the aedeagus described (Allen and Hutton Reference Allen and Hutton1969; Wittmer Reference Wittmer1976; Roza et al. Reference Roza, Constantin and Mermudes2019; López-Pérez and Zaragoza-Caballero Reference López-Pérez and Zaragoza-Caballero2021; Ferreira Reference Ferreira2025). It consists of a large phallobase that extends past the base of the parameres and median lobe; the parameres are fused basally, articulating with the median lobe, and have a subbasal medial arch that encircles the median lobe and extends almost to the parameres’ apex, which are spatulate with rounded apical margin, ventrally granulate, and bear sparse setae; and the median lobe has the base dorsally oriented (where the parameres articulate), is slightly tapered towards the apex in dorsal view, and is curved from the base to the apex in lateral view, with the apex dorsally oriented (Fig. 6).
Some variability in the paramere apex shape and the median lobe curvature is seen but seems to be species-level variation rather than generic characteristics. Although López-Pérez and Zaragoza-Caballero (Reference López-Pérez and Zaragoza-Caballero2021) use the parameres’ apex to distinguish Platyphalus and Stenodrilus, the difference is subtle. Moreover, it is hard to know the effect of any scanning electron microscope preparation on the parameres, which are thin and fragile. The structure preparation for scanning electron microscope imaging, however, is not described in López-Pérez and Zaragoza-Caballero (Reference López-Pérez and Zaragoza-Caballero2021). Further investigation with the dissection of more specimens in both genera is necessary to determine the true extent of the genitalic variation and whether it is a suitable character for generic diagnosis.
Acknowledgements
I thank Christoffer Fägerström and Jadranka Rota (MZLU) and François Génier and Robert Anderson (CMN) for their assistance and the loan of the specimens examined in this study. I also thank Vinicius S. Ferreira and two anonymous reviewers for corrections and suggestions that improved the manuscript. I acknowledge the Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) for funding (proc. 205.818/2022 and 205.819/2022) and for the photographic system acquired through a grant (proc. 110.040/2014).
Competing interests
The author declares that he has no competing interests.
