Research Article |
Mariana Raquel Chani-Posse‡, Maryzender Erceliz Rodríguez-Melgarejo‡§|, José Manuel Ramírez-Salamanca‡¶
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Corresponding author: Mariana Raquel Chani-Posse ( mchani@mendoza-conicet.gob.ar ) Academic editor: Vinicius S. Ferreira
© 2025 Mariana Raquel Chani-Posse, Maryzender Erceliz Rodríguez-Melgarejo, José Manuel Ramírez-Salamanca.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Abstract
Konradus Chani-Posse & Ramírez-Salamanca, 2020 was originally described as a monotypic genus from the tropical Andes of Ecuador. However, a re-examination of type material from species previously classified under the genus Philonthus Stephens, 1829, along with additional, previously unstudied specimens collected from high-altitude regions (above 2000 m) in the Andes of Ecuador, Peru, and Bolivia, prompted a more comprehensive reassessment of the genus. This study presents a revision of Konradus and introduces Yuracarus gen. nov., based on external morphology and male and female sexual characters. Two species previously considered “false Philonthus” (Ph. actinus Bernhauer, 1917 and Ph. diamantinus Bernhauer, 1917) are transferred to Konradus and Yuracarus gen. nov., respectively: K. actinus (Bernhauer), new comb., and Y. diamantinus (Bernhauer), new comb. Additionally, two species from Peru are newly described within Konradus: K. cuscensis sp. nov. and K. trescrucensis sp. nov., while three new species are described within Yuracarus: Y. cosnipatensis sp. nov., Y. napoensis sp. nov., and Y. yunguensis sp. nov. A key to species, diagnoses, descriptions and/or redescriptions, illustrations, a distributional map, and a phylogenetic analysis are included. Lectotypes are designated for Philonthus actinus Bernhauer and Philonthus diamantinus Bernhauer. Cladistic analysis confirms the monophyly of Konradus and Yuracarus within the Andean clade (AC) of Neotropical Philonthina and supports Yuracarus as a distinct genus. Both genera are part of a well-supported clade characterized by the presence of sub-bilobed and subtriangular protarsomeres 2 and 3, bearing discal setae on their ventral surface, as well as a horseshoe-shaped accessory sclerite associated with female tergum 10.
Keywords
Bolivia, Ecuador, morphology, new genus, Peru, Philonthina, Staphylininae, Tropical Andes
1. Introduction
Previous studies on Philonthina have consistently demonstrated the non-monophyly of the megadiverse and globally distributed genus Philonthus Stephens, 1829 (e.g.,
In addition to revising Konradus, describing the new genus Yuracarus, and reclassifying and redescribing Ph. actinus and Ph. diamantinus, we describe five new species based on recently examined material collected from high-altitude regions in the Andes of Ecuador, Peru, and Bolivia. Of these, two species are assigned to Konradus, and three to the newly established genus Yuracarus, which also includes Ph. diamantinus. Finally, we test the monophyly of both genera and explore their phylogenetic relationships within the Neotropical lineage of Philonthina.
2. Material and methods
2.1. Examination of material and terminology
The material studied was borrowed from the following depositories:
Beetle specimens were examined using a Nikon SMZ 745 dissecting microscope. They were mostly examined as pinned dry specimens, but two of them were first relaxed in warm soapy water, rinsed, disarticulated and examined as wet preparations in ethanol. Techniques for the preparation and examination of male and female genitalia follow
HW Head capsule maximum width (measured at widest point);
HL Length of head capsule, from anterior margin of frontoclypeus to neck constriction (along midline);
Lp2L, Lp3L Length of second or third labial palpomere;
PW Pronotum maximum width;
PL Pronotum length along midline;
EL Eye length (seen from above);
TL Temple length (from the posterior margin of the eye to the nuchal groove; seen from above);
NW Neck width;
S1, S5 Length of first or fifth (last) metatarsomere;
EtL Elytron length at side (straight line from humerus to apex; seen from above).
Terminology follows authors and criteria as stated in
During the preparation of this work, ChatGPT Open AI was used solely in order to improve language and readability. After this tool/service was used, the content was reviewed and edited as needed, and full responsibility for the content of the publication is taken by the authors herein.
2.2. Phylogenetic analysis
2.2.1. Outgroup and ingroup taxa
A total of twenty-three terminal units are included in the phylogenetic analysis. In addition to Konradus leehermani, the type species, this analysis includes the other three species recognized as Konradus, along with the four species belonging to the newly established genus Yuracarus gen. nov. Fifteen additional species are also included, representing genera (including type species* or species within the type species group) from Philonthina and its NL, with a focus on the Andean clade (AC hereafter) (
2.2.2. Characters
Sixty-seven morphological characters (see File S1) were coded and scored for the 23 terminal units, with their selection and definition following
1* Body, metallic coloration: [0] absent or limited to elytra; [1] limited to head, thorax and elytra (e.g., Fig.
2 Antennal insertions (ai), position in relation to frontoclypeus and eye (see
3 Antennal insertions (ai), position (see
4 Antennae, antennomeres 4–10, apical margin, shape: [0] concave to truncate; [1] distinctly convex.
5 Antennae, antennomere 5, shape: [0] elongate; [1] quadrate.
(6) Sexual dimorphism, frontoclypeus, emargination: [0] absent (i.e., both male and female without emargination); [1] present (emarginate in males) (
7 Eyes, relative length to that of tempora: [0] subequal; [1] eyes distinctly longer than tempora; [2] eyes distinctly shorter than tempora.
8* Head, eye shape: [0] not protruding; [1] protruding (Fig.
9 Head shape, maximum width: [0] close to or at its anterior margin; [1] at its halfway between the anterior and posterior margin; [2] at or close to the posterior margin.
(10) Head, anterior angles next to eye margin: [0] without a distinct border; [1] with a distinct border (
11* Head, posterior angles: [0] almost obsolete (Fig.
12 Head, postmandibular ridge: [0] absent; [1] present (Fig.
13 Head, distance between antennal insertions relative to shortest distance between anterior margin of eye to antennal insertion: [0] 2× or more; [1] distinctly less than 2×.
14 Head, labrum shape: [0] transverse; [1] subconical (
15 Head, ventral surface, microsculpture: [0] distinct (i.e., punctate near gular sutures); [1] not distinct.
16 Gular sutures, development from anterior margin of mentum to base of head: [0] not joined; [1] joined halfway; [2] joined rather close to base of head.
17 Labial palpus, relative length of palpomeres 3 (p3, apical) and 2 (p2, preapical) (p3/p2): [0] p3 distinctly shorter than p2; [1] p3 and p2 subequal in length; [2] p3 distinctly longer than p2.
18 Maxillary palpus, relative length of palpomeres 3 (p3, preapical) and 2 (p2) (p3/p2): [0] p2 not distinctly longer than p3 (maximum a3/a2 ratio 1.2); [1] p2 distinctly longer than p3.
(19) Maxillary palpus, relative length of palpomeres 4 (p4, apical) and 3 (p3, preapical) (p4/p3): [0] 1.5 at most; [1] longer than 1.5.
20 Maxillary palpus, relative width of palpomeres 4 (p4, apical) and 3 (p3, preapical) (p4/p3): [0] p4 not distinctly narrower than p3 for most of its length; [1] p4 distinctly narrower than p3 for most of its length.
(21) Maxillary palpus, palpomere 4 (apical), shape: [0] subacute, i.e., narrowed at base and evenly converging towards apex; [1] fusiform to expanded apically, i.e., narrowed at base but not (or not evenly) converging towards apex (
22 Prothorax, pronotum shape: [0] broadened posteriad; [1] parallel-sided or broadened at the anterior angles; [2] pronotum broadened anteriad behind the anterior angles.
23* Prothorax, pronotum, lateral margins: [0] straight and/or arcuate, not constricted before reaching posterior angles (Fig.
(24) Prothorax, pronotum, anterior angles: [0] not extended anteriad; [1] extended anteriad (
25 Prothorax, pronotum, posterior angles: [0] rounded to slightly obtuse (
(26) Prothorax, pronotum, deflection, ventral view: [0] absent; [1] present.
27 Prothorax, large lateral setiferous puncture (llsp), position in relation to superior marginal line (sml) of pronotum (see
(28) Prothorax, hypomeron, inferior marginal line (iml), development (see
29 Prothorax, prosternum, shape of longitudinal carina: [0] forming well-defined, sharp ridge at least in its basal part; [1] more or less rounded, from obtuse ridge to smooth longitudinal prominence of prosternum.
30 Male protarsi, tarsomeres 2–3: [0] not bilobed (posterior margin moderately curved to straight); [1] sub-bilobed (Fig.
31 Male protarsi, tarsomeres 2–3, shape: [0] rectangular (i.e., flattened and rather transverse); [1] more or less cylindrical; [2] subtriangular (i.e., flattened and widened distally).
32 Protarsomeres 2–3, ventral surface, discal (adhesive) setae: [0] absent; [1] present.
33* Protarsomeres 2–3, ventral surface, discal setae (if present), distribution: [0] rather few, short and scattered (i.e., not forming a patch covering next segment) (Fig.
(34) Protarsomeres 2–3, ventral surface, discal setae (if present), shaft, microsculpture (see Chani-Posse and Ramírez-Salamanca 2020): [0] smooth (fig. 3C); [1] striate (fig. 3H).
(35) Protarsomeres 2–3, ventral surface, discal setae (if present), shaft, shape (see Chani-Posse and Ramírez-Salamanca 2020): [0] cylindrical, distinctly broadened apically (fig. 3B); [1] subcylindrical, gradually narrowed apically (fig. 3F).
36 Protarsomeres 2–3, ventral surface, marginal setae (see Chani-Posse and Ramírez-Salamanca 2020): [0] absent (if present only 1–2 fine setae at each lateroposterior angle) (fig. 2I, N, O); [1] present (e.g., fig. 2J, K, Q).
37 Protarsomeres 2–3, ventral surface, marginal setae, distribution (if present) (see Chani-Posse and Ramírez-Salamanca 2020): [0] posteriad (fig. 2Q); [1] lateral (fig. 2T).
38 Sexual dimorphism, profemur, shape: [0] absent (i.e., equally wide in both sexes); [1] present (i.e., wider in males).
39* Male profemora, shape: [0] not angulate medially; [1] distinctly angulate medially (Fig.
(40) Male profemora, spines, all along lateral face: [0] absent; [1] present.
41 Mesoventrite, intercoxal ridge (see
42* Mesotarsomere 2, ventral surface, discal setae: [0] absent and if so, barely visible; [1] present, distinct (Fig.
43* Elytral punctation: [0] slightly punctate (i.e., distance between punctures exceeding 2× diameter of puncture) (e.g., Figs
(44) Elytra, surface shape: [0] distinctly flat; [1] with a couple of swellings.
45 Elytra, middle suture, elevation: [0] absent; [1] present.
(46) Abdomen, tergum 3, posterior basal transverse carina (PBTC): [0] absent; [1] present.
(47) Abdomen, tergum 4, PBTC: [0] absent; [1] present.
(48) Abdomen, tergum 5, PBTC: [0] absent; [1] present.
49 Abdomen, tergum 6, PBTC: [0] absent; [1] present.
50 Abdomen, sternum 3, basal transverse carina, shape: [0] arcuate; [1] acutely pointed to subangulate medially.
51 Abdomen, terga 3–7, setae: [0] almost glabrous; [1] scarcely setose (i.e., setae sparse and posteriad); [2] densely setose.
52 Abdomen, terga 3–5, rugose microsculpture, posterior basal transverse carina: [0] absent; [1] present.
53* Abdomen, tergum 8, hind margin: [0] not angulate medially (Fig.
(54) Male sternum 8, apical margin, notch (see
55 Male sternum 8, apical margin, emargination: [0] slight; [1] moderate; [2] deep.
(56) Male sternum 9, distal portion, emargination (if distinct): [0] acute; [1] subangulate to concave; [2] slightly arcuate.
57 Male sternum 9, basal portion, left end: [0] acute; [1] angulate; [2] rounded; [3] subacute.
58 Male tergum 10, shape: [0] emarginate medio-apically; [1] apically subtruncate to wide and subangulate or arcuate; [2] concave medio-apically to apically truncate; [3] apically distinctly subacute to acute.
59 Aedeagus, paramere(s), sensory peg setae: [0] absent; [1] present.
60 Aedeagus, paramere(s), degree of attachment to median lobe: [0] fused to median lobe only at base, otherwise paramere(s) distinctly separated from median lobe along entire length; [1] fused entirely to median lobe.
61 Aedeagus, paramere(s), shape: [0] elongate but rather flattened; [1] plate-like; [2] rod-like.
(62) Aedeagus, median lobe, apical half, lateral view (see
63 Aedeagus, median lobe, apical fourth, dorsal view: [0] gradually narrowed; [1] distinctly narrowed.
64 Aedeagus, median lobe, apex, dorsal view: [0] acute; [1] subacute to rounded.
65 Female tergum 10, shape: [0] emarginate medio-apically; [1] apically subtruncate to wide and subangulate or arcuate; [2] concave medio-apically to apically truncate; [3] apically distinctly subacute to acute.
66 Female abdomen, genitalia, accessory sclerite: [0] absent; [1] present.
67* Female abdomen, genitalia, accessory sclerite, shape (if present): [0] horseshoe-shaped (Fig.
2.2.3. Procedure
Matrix analysis was performed using Mesquite v3.81 (
Bayesian inference was conducted with MrBayes v. 3.2.7 (
In MP analysis, nodes with BS ≥ 2 were considered well-supported, while those with BS = 1 were regarded as weakly supported. For ML, nodes with SH-aLRT ≥ 80 and UFB ≥ 95 were considered well-supported, nodes with either SH-aLRT < 80 or UFB < 95 as weakly supported, and nodes with both SH-aLRT < 80 and UFB < 95 as unsupported. In BI analysis, nodes with Bayesian posterior probability (PP) ≥ 0.90 were deemed well-supported, PP values of 0.80–0.89 as weakly supported, and PP < 0.80 as unsupported. Nodes that were unsupported were typically not discussed when comparing topologies between methods or addressing monophyly.
All trees from both ML and BI analyses were visualized using FigTree v1.4.5_pre (
3. Results
3.1. Taxonomy
Konradus
Konradus
Chani-Posse and Ramírez-Salamanca 2020: 239;
Species composition.
Four species: Konradus actinus (Bernhauer) comb. nov. (Bolivia); Konradus cuscensis sp. nov. (Peru); Konradus leehermani Chani-Posse & Ramírez-Salamanca (Ecuador); Konradus trescrucensis sp. nov. (Peru).
Type species.
Konradus leehermani Chani-Posse & Ramírez-Salamanca, 2020.
Diagnosis and description.
As in
Recognition.
Konradus can be easily recognized among other Neotropical genera of Philonthina by its distinct and entirely metallic-colored habitus, characterized by an elongate head and pronotum, with posterior angles of the head almost obsolete, gular sutures joined posteriorly before the neck, eyes distinctly shorter than the tempora, and protarsomeres 1–4 subequal in length, slightly to moderately widened apically, bearing both discal and marginal striate setae on the ventral surface, with the discal setae sparse and not forming a patch. Furthermore, Konradus can be distinguished from Yuracarus gen. nov. by several key features. The head of Konradus has posterior angles that are almost obsolete, while those of Yuracarus are distinctly obtuse. The profemur in Konradus is subcylindrical and not angulate medially, whereas in Yuracarus it is distinctly angulate in both sexes. Additionally, the protarsomeres 1–4 of Konradus have discal setae that are scattered and do not form a patch, in contrast to those of Yuracarus, where the discal setae form a patch on the ventral surface.
Distribution.
Konradus is currently known from the Tropical Andes of Ecuador, with new records from Peru and Bolivia (Fig.
Key to species of Konradus Chani-Posse & Ramírez-Salamanca, 2020
| 1 | Head about as wide as pronotum (HW/PW = 1.0–1.1), pronotum parallel-sided, not narrowed posteriad (e.g. Fig. |
2 |
| – | Head wider than pronotum (HW/PW = 1.2), pronotum narrowed posteriad (Fig. |
Konradus leehermani Chani-Posse & Ramírez-Salamanca |
| 2 | Antennomeres 7–8 elongate or quadrate, distance separating medial interocular punctures on frons at least 2.5× distance separating medial punctures from lateral punctures | 3 |
| – | Antennomeres 7–8 transverse, distance separating medial interocular punctures on frons not more than 2× distance separating medial punctures from lateral punctures | Konradus trescrucensis sp. nov. |
| 3 | Antennomeres 7–8 elongate, distance separating medial interocular punctures on frons 2.5× distance separating medial punctures from lateral punctures | Konradus actinus (Bernhauer), new comb. |
| – | Antennomeres 7–8 quadrate, distance separating medial interocular punctures on frons 3× distance separating medial punctures from lateral punctures | Konradus cuscensis sp. nov. |
Konradus leehermani
Konradus leehermani
Chani-Posse & Ramírez-Salamanca, 2020: 241–243 (description, figures, distribution, phylogeny);
Diagnosis.
Konradus leehermani can be identified among other species of Konradus by the following combination of characters: antennomeres 7–8 elongate, the distance separating the medial interocular punctures on the frons not more than twice the distance separating the medial punctures from the lateral punctures, the head wider than the pronotum at its widest point, and the pronotum narrowed posteriad.
A Konradus leehermani Chani-Posse & Ramírez-Salamanca [
Philonthus diamantinus Bernhauer [
Description and distribution.
As in Chani-Posse and Ramírez-Salamanca (2020).
Konradus actinus , comb. nov.
Philonthus actinus
Type material.
Lectotype ♂, here designated (Fig.
Konradus actinus (Bernhauer): A male sternum 8; B sternum 9; C female tergum 8; D female tergum 10. Konradus trescrucensis sp. nov.: E sternum 9. Konradus cuscensis sp. nov: F male tergum 8; G sternum 9; H male tergum 10; I gonocoxites of female genital segment; J accessory sclerite. Yuracarus napoensis sp. nov.: K male sternum 8; L sternum 9. Yuracarus yunguensis sp. nov.: M sternum 9; N male tergum 10; O gonocoxites of female genital segment; P female tergum 10. Yuracarus cosnipatensis: Q female tergum 8. Yuracarus diamantinus (Bernhauer): R female tergum 10; S gonocoxites of female genital segment; T accessory sclerite. Scale bars: 0.5 mm (A–I, K–S); 0.2 mm (J, T).
Additional material examined.
(2♂, 2♀). BOLIVIA: Cochabamba, 70 km NE Cochabamba, 4.IV.1978, L.& C.W. O’ Brien & G.B. Marshall, 1♂ (
Aedeagus, dorsal and lateral views: A Konradus actinus (Bernhauer), B Konradus cuscensis sp. nov., C Konradus trescrucensis sp. nov., D Yuracarus diamantinus (Bernhauer), E Yuracarus yunguensis sp. nov., F Yuracarus cosnipatensis sp. nov., G Yuracarus napoensis sp. nov. Scale bars: 0.5 mm.
Diagnosis.
Konradus actinus can be identified among other species of Konradus by the following combination of characters: antennomeres 7–8 elongate, the distance separating the medial interocular punctures on the frons 2.5× distance separating the medial punctures from the lateral punctures, the head as wide as pronotum at its widest point, and the pronotum parallel-sided.
Redescription.
Body length 9.4–9.8 mm (5.4–5.6 mm, abdomen excluded), body overall scarcely setose, almost glabrous. — Colouration: body metallic green, with last abdominal segments metallic purple to greenish purple. — Head: moderately to slightly longer than wide (HW/HL= 0.8–0.9), as wide as pronotum at widest point (HW/ PW= 1.0); epicranium with medial interocular punctures separated by distance 2.5× as large as distance separating medial punctures from lateral punctures. Eyes distinctly shorter than temples (EL/TL = 0.5–0.7) seen from above. Antennae with first antennomere distinctly shorter than second and third combined, third about 1.2–1.4× as long as second, fourth-eighth longer than wide, ninth-tenth about as long as wide. Labial palpus with third palpomere 1.5× as long as second. Maxillary palpus with fourth palpomere (apical) 1.6–1.8× as long as third. Neck about 0.4–0.5× as wide as head at widest point. — Pronotum: distinctly to moderately longer than wide (PW/PL = 0.7–0.8), dorsal rows of punctures each with 5 punctures. Prosternum without distinct mid-longitudinal carina. — Legs: Mesotarsi about as long as mesotibiae, metatarsi about as long as metatibiae, first metatarsomere shorter than fifth (S1/S5= 0.7–0.8). — Elytra: at sides slightly longer than pronotum at midline (EtL/PL=1.1). — Male sexual characters: Sternum 8 slightly emarginate medio-apically, emargination without semi-membranous extension (Fig.
Konradus cuscensis sp. nov.
Type material.
Holotype ♂ (Fig.
Diagnosis.
Konradus cuscensis can be identified among other species of Konradus by the following combination of characters: antennomeres 7–8 quadrate, the distance separating the medial interocular punctures on the frons at least 3.0× distance separating the medial punctures from the lateral punctures, the head as wide as pronotum at its widest point, and the pronotum parallel-sided.
Description.
Body length 8.5–9.8 mm (5.2–5.9 mm, abdomen excluded), body overall scarcely setose, almost glabrous. — Colouration: entire body varying from metallic purple to greenish purple. — Head: distinctly longer than wide (HW/HL= 0.7), as wide as pronotum at widest point (HW/ PW= 1.0); epicranium with medial interocular punctures separated by distance 3× as large as distance separating medial punctures from lateral punctures. Eyes distinctly shorter than temples (EL/TL = 0.6) seen from above. Antennae with first antennomere distinctly shorter than second and third combined, third about 1.2× as long as second, fourth-sixth longer than wide, seventh-tenth about as long as wide. Labial palpus with third palpomere 1.5× as long as second. Maxillary palpus with fourth palpomere (apical) 1.7× as long as third. Neck about 0.6× as wide as head at widest point. — Pronotum: distinctly longer than wide (PW/PL = 0.7), dorsal rows of punctures each with 5 punctures. Prosternum without distinct mid-longitudinal carina. — Legs: Mesotarsi slightly shorter than mesotibiae, metatarsi about as long as metatibiae, first metatarsomere shorter than fifth (S1/S5= 0.9). — Elytra: at sides as long as pronotum at midline (EtL/PL=1.0). — Male sexual characters: Sternum 8 slightly emarginate medio-apically, emargination without semi-membranous extension. Aedeagus with median lobe distinctly narrowed apically into rather blunt, subacute apex, internal sac with one pair of elongate and sclerotized structures (Fig.
Phylogenetic reconstructions indicating relationships of Konradus Chani-Posse & Ramírez-Salamanca and Yuracarus gen. nov. within the Andean clade: A most parsimonious tree with black circles as exclusive (non-homoplastic) synapomorphies and white circles as non-exclusive (homoplastic) synapomorphies. Numbers in bold below the branches are absolute Bremer support values; B maximum Likelihood tree with support values SH-aLRT / UFB; C fifty percent majority consensus tree with posterior probabilities.
Geographical distribution.
Konradus cuscensis sp. nov. is currently recorded only from the locality of Pillahuata (Cosñipata district, Paucartambo province) from Cuzco department, in the Tropical Andes of Peru, at elevations beyond 2000 m (Fig.
Etymology.
The specific name cuscensis is a Latinized adjective referring to the locality of Cuzco, where this species was collected.
Konradus trescrucensis sp. nov.
Type material.
Holotype ♂, with labels: [white printed label] “PERU: CU, Tres Cruces / Abra Acjanaco, Bosque / Maderero, 3600m / ca. 13°18’S/ 71°40’W / 12.vii.1991 D. Silva”, “Muestreo / por Golpeo”, [red label] “Holotype Konradus / trescrucensis Chani-Posse / et al. 2025” (
Diagnosis.
Konradus trescrucensis can be identified among other species of Konradus by the following combination of characters: antennomeres 7–8 transverse, the distance separating the medial interocular punctures on the frons 2.0× distance separating the medial punctures from the lateral punctures, the head not distinctly wider than pronotum at its widest point, and the pronotum parallel-sided.
Description.
Body length 8.5–8.8 mm (4.9–5.1 mm, abdomen excluded), body overall scarcely setose, almost glabrous. — Colouration: head, thorax, elytra, and abdominal segments 6 and 7 metallic purple to purplish-green; scutellum and rest of abdomen metallic green. — Head: slightly longer than wide (HW/HL= 0.9), as wide as to slightly wider than pronotum at widest point (HW/ PW= 1.0–1.1); epicranium with medial interocular punctures separated by distance 2.0× as large as distance separating medial punctures from lateral punctures. Eyes distinctly shorter than temples (EL/TL = 0.6) seen from above. Antennae with first antennomere moderately to slightly shorter than second and third combined, third about 1.2–1.4× as long as second, fourth and fifth longer than wide, sixth about as long as wide, seventh-tenth transverse. Labial palpus with third palpomere 1.5× as long as second. Maxillary palpus with fourth palpomere (apical) 2.0× as long as third. Neck about 0.4× as wide as head at widest point. — Pronotum: distinctly to moderately longer than wide (PW/PL = 0.7–0.8), dorsal rows of punctures each with 5 punctures. Prosternum without distinct mid-longitudinal carina. — Legs: Mesotarsi slightly shorter than mesotibiae, metatarsi slightly shorter than metatibiae, first metatarsomere shorter than fifth (S1/S5= 0.8). — Elytra: at sides as long as to slightly longer than pronotum at midline (EtL/PL=1.0–1.1). — Male sexual characters: Sternum 8 slightly emarginate medio-apically, emargination without semi-membranous extension. Aedeagus with median lobe gradually narrowed from base into a subacute apex, internal sac with one pair of elongate and sclerotized structures (Fig.
Geographical distribution.
Konradus trescrucensis sp. nov. is currently recorded only from the locality of Tres Cruces (Cosñipata district, Paucartambo province) from Cuzco department, in the Tropical Andes of Peru, at 3600 m of altitude (Fig.
Etymology.
The specific name trescrucensis is a Latinized adjective referring to the locality of Tres Cruces, where this species was collected.
Yuracarus gen. n.
Species composition (four species):
Yuracarus cosnipatensis sp. nov. (Peru); Yuracarus diamantinus (Bernhauer) comb. nov. (Bolivia); Yuracarus napoensis sp. nov. (Ecuador); Yuracarus yunguensis sp. nov. (Bolivia, Peru)
Type species.
Philonthus diamantinus Bernhauer, 1917, here designated.
Diagnosis.
The most distinctive characters of Yuracarus are its slender body shape, as well as its predominantly metallic-colored habitus (excluding the abdomen), in addition to the profemur distinctly angulate medially, the protarsomeres 1–4 enlarged, sub-bilobed, and rather flattened, with a patch of striate and petiolate discal setae, also present on the meso- and metatarsomeres, although less distinct due to the meso- and metatarsomeres being distinctly narrower than the protarsomeres, as well as the hind margin of tergum 8 truncate and angulate medially in both sexes. In addition to these characters, it differs from other Neotropical genera with enlarged protarsomeres and discal setae currently known (i.e., Delgadobius, Konradus, Inesius, Rhaegalius and Atopocentrum) by the gular sutures being joined anteriorly, whereas in Delgadobius, Konradus, Inesius, and Rhaegalius, the gular sutures are either joined posteriorly (Konradus, Inesius, Rhaegalius and Atopocentrum) or not joined (Delgadobius).
Description.
Length 8.8–10.7 mm. Body elongate, more or less parallel sided, slightly widening toward elytra and tapering toward sixth visible abdominal segment, scarcely punctate (Fig.
Distribution.
Yuracarus is currently known from the Tropical Andes of Ecuador, Peru and Bolivia (Fig.
Etymology.
Yuracarus is a masculine, Latinized generic name derived from Yuracaris, the type locality of this new genus. The root “Yura-” is combined with the Latin suffix “-carus”, denoting its association with the locality.
Key to species of Yuracarus gen. nov.
| 1 | Antennomeres 8–10 quadrate, eyes at least 0.6× the length of temples | 2 |
| – | Antennomeres 8–10 elongate, eyes not longer than half the length of temples | Yuracarus diamantinus (Bernhauer), new comb. |
| 2 | Distance separating medial interocular punctures on frons at least 2.5× distance separating medial punctures from lateral punctures | 3 |
| – | Distance separating medial interocular punctures on frons not more than 2.0× distance separating medial punctures from lateral punctures | Yuracarus yunguensis sp. nov. |
| 3 | Head narrower than pronotum (HW/ PW = 0.9), distance separating medial interocular punctures on frons 3.0× distance separating medial punctures from lateral punctures | Yuracarus napoensis sp. nov. |
| – | Head as wide as pronotum (HW/ PW = 1.0), distance separating medial interocular punctures on frons 2.5× distance separating medial punctures from lateral punctures | Yuracarus cosnipatensis sp. nov. |
Yuracarus cosnipatensis sp. nov.
Type material.
Holotype ♂, with labels: [printed white label] “PERU: CU, Kosñipata, Wayqecha, / Wayqecha Trocha / Canopy 13°11’32.8’’S, / 71°35’16.0’’W, 2931 m”, “02.xi.2017, cloud forest, / sifting of leaf litter, M. / Rodríguez & L. Pérez leg”, [red label] “Holotype Yuracarus / cosnipatensis Chani-Posse / et al. 2025” (
Diagnosis.
Yuracarus cosnipatensis can be identified among other species of Yuracarus by the following combination of characters: antennomeres 8–10 quadrate, eyes at least 0.6× the length of the temples, the distance separating the medial interocular punctures on the frons at least 2.5× distance separating the medial punctures from the lateral punctures, and the head as wide as the pronotum (HW/PW = 1.0).
Description.
Body length 8.8–9.0 mm (4.8–5.0 mm, abdomen excluded), body overall scarcely setose, almost glabrous. — Colouration: Head and thorax dark metallic green, elytra goldish metallic green, abdomen shiny black to piceous-black; antennae, palpi and legs piceous to piceous-black. — Head: about as long as wide (HW/HL= 0.9–1.0), as wide as pronotum at widest point (HW/ PW= 1.0); epicranium with medial interocular punctures separated by distance 2.5× as large as distance separating medial punctures from lateral punctures. Eyes distinctly shorter than temples (EL/TL = 0.7) seen from above. Antennae with first antennomere moderately to distinctly shorter than second and third combined, third 1.2–1.3× as long as second, fourth-seventh longer than wide, eighth-tenth about as long as wide. Labial palpus with third palpomere 1.5× as long as second. Maxillary palpus with fourth palpomere (apical) 1.5× as long as third. Neck about 0.4–0.5x as wide as head at widest point. — Pronotum: moderately to slightly longer than wide (PW/PL = 0.8–0.9), dorsal rows of punctures each with 4 punctures. Legs: Mesotarsi as long as mesotibiae, metatarsi slightly shorter than metatibiae, first metatarsomere shorter than fifth (S1/S5= 0.8). — Elytra: at sides distinctly longer than pronotum at midline (EtL/PL=1.3). — Male sexual characters: Sternum 8 deeply emarginate medio-apically. Aedeagus with median lobe distinctly narrowed at its apical fourth into a subacute apex, internal sac with one pair of elongate and sclerotized structures (Fig.
Geographical distribution.
Yuracarus cosnipatensis sp. nov. is currently recorded only from the locality of Cosñipata (Cosñipata district, Paucartambo province) in Cuzco department, from the Tropical Andes of Peru, at 2837 and 2931 m of altitude (Figs
Etymology.
The specific name cosnipatensis is a Latinized adjective referring to the locality of Cosñipata, where this species was collected.
Yuracarus diamantinus , comb. nov.
Philonthus diamantinus
Type material.
Lectotype ♂, here designated (Figs
Additional material examined.
BOLIVIA: Chapare, Yungas, 29–31.I.76, Bolle, 2900m, ♂ (
Diagnosis.
Yuracarus diamantinus can be identified among other species of Yuracarus by the following combination of characters: antennomeres 8–10 elongate, eyes not longer than half the length of the temples, and the distance separating the medial interocular punctures on the frons not more than 2.0× distance separating the medial punctures from the lateral punctures.
Redescription.
Body length 10.2–10.3 mm (5.0–5.2 mm, abdomen excluded), body overall scarcely setose, almost glabrous (Fig.
Geographical distribution.
Yuracarus diamantinus is currently recorded only from the Andes of Bolivia, at elevations ranging from approximately 2000 m to 2900 m (Fig.
Yuracarus napoensis sp. nov.
Type material.
Holotype ♂, with labels: [white printed label] “ECU. Napo, 10200’ / Papallacta / 21.V.82, H. Frania / canopy dead tree”, [red label] “Holotype Yuracarus / napoensis Chani-Posse et al. 2025” (
Diagnosis.
Yuracarus napoensis can be identified among other species of Yuracarus by the following combination of characters: antennomeres 8–10 quadrate, eyes at least 0.6× the length of the temples, the distance separating the medial interocular punctures on the frons 3× distance separating the medial punctures from the lateral punctures, and the head narrower than pronotum (HW/PW = 0.9).
Description.
Body length 9.6–10.7 mm (5.1–5.7 mm, abdomen excluded), body overall scarcely setose, almost glabrous. — Colouration: Head and thorax metallic greenish blue, elytra mostly metallic purple to greenish purple apically, abdomen shiny black to piceous-black; antennae, palpi and legs piceous to piceous-black. — Head: about as long as wide (HW/HL= 1.0), slightly narrower than pronotum at widest point (HW/ PW= 0.9); epicranium with medial interocular punctures separated by distance 3× as large as distance separating medial punctures from lateral punctures. Eyes distinctly shorter than temples (EL/TL = 0.6) seen from above. Antennae with first antennomere distinctly shorter than second and third combined, third 1.1× slightly longer than second, fourth-seventh longer than wide, eighth-tenth about as long as wide. Labial palpus with third palpomere 1.5× as long as second. Maxillary palpus with fourth palpomere (apical) 1.8× as long as third. Neck about 0.4× as wide as head at widest point. — Pronotum: moderately longer than wide (PW/PL = 0.8), dorsal rows of punctures each with 4 punctures. — Legs: Mesotarsi shorter than mesotibiae, metatarsi shorter than metatibiae, first metatarsomere shorter than fifth (S1/S5= 0.8). — Elytra: at sides moderately longer than pronotum at midline (EtL/PL=1.2). — Male sexual characters: Sternum 8 moderately emarginate medio-apically (Fig.
Geographical distribution.
Yuracarus napoensis sp. nov. is currently recorded only from the locality of Napo in the Andes of Ecuador, at approximately 2370 and 3100 m of altitude (Fig.
Etymology.
The specific name napoensis is a Latinized adjective referring to the locality of Napo, where this species was collected.
Yuracarus yunguensis sp. nov.
Type material.
Holotype ♂, with labels: [white label] “BOLIVIA: La Paz: / Sorata, 2800m / XI.1984, L. E. Peña, leg. / FIELD MUS. NAT. HIST.”, [red label] “Holotype Yuracarus / yunguensis Chani-Posse / et al. 2025” (
Diagnosis.
Yuracarus yunguensis can be identified among other species of Yuracarus by the following combination of characters: antennomeres 8–10 quadrate, eyes at least 0.6× the length of the temples, and the distance separating the medial interocular punctures on the frons not more than 2.0× distance separating the medial punctures from the lateral punctures.
Description.
Body length 9.2–10.7 mm (5.2–5.6 mm, abdomen excluded), body overall scarcely setose, almost glabrous. — Colouration: Head and thorax metallic greenish blue, elytra mostly metallic purple basally to greenish blue apically, abdomen shiny black to piceous-black; antennae, palpi and legs piceous to piceous-black. — Head: slightly longer than (HW/HL= 0.9), as long as pronotum at widest point (HW/ PW= 1.0); epicranium with medial interocular punctures separated by distance 2.0× as large as distance separating medial punctures from lateral punctures. Eyes moderately shorter than temples (EL/TL = 0.8) seen from above. Antennae with first antennomere distinctly shorter than second and third combined, third 1.1× slightly longer than second, fourth-seventh longer than wide, eighth-tenth about as long as wide. Labial palpus with third palpomere 1.5× as long as second. Maxillary palpus with fourth palpomere (apical) 1.2× as long as third. Neck about 0.4× as wide as head at widest point. — Pronotum: moderately longer than wide (PW/PL = 0.8), dorsal rows of punctures each with 4 punctures. — Legs: Mesotarsi shorter than mesotibiae, metatarsi shorter than metatibiae, first metatarsomere slightly shorter than fifth (S1/S5= 0.9). — Elytra: at sides distinctly longer than pronotum at midline (EtL/PL=1.3). — Male sexual characters: Sternum 8 deeply emarginate medio-apically. Aedeagus with median lobe gradually narrowed from base into rather acute apex, internal sac with one pair of elongate and sclerotized structures (Fig.
Geographical distribution.
Yuracarus yunguensis sp. nov. is currently recorded from the Andes of Peru and Bolivia, at 2800 m and 4100 m of altitude (Fig.
Etymology.
The specific name yunguensis is a Latinized adjective referring to the Yungas ecoregion, where this species was collected.
3.2. Phylogeny
The analysis of the data matrix (File S1) produced one most parsimonious tree (MPT) (Fig.
Of the 21 nodes recovered by MP, 20 were supported by one, two, or all three inference methods. MP supported 20 nodes, while ML and BI each supported 10 nodes, respectively. The single MPT, along with the trees obtained by BI and ML and their support values, are shown in Fig.
Overall, the consensus topologies recovered by MP, BI, and ML analyses were consistent at the generic level, showing that Konradus and Yuracarus each form distinct clades within a weakly supported AC only by MP (BS = 1). Incongruences mainly arose between MP and the model-based methods (ML and BI) in resolving basal relationships within the AC, with both ML and BI failing to resolve these relationships, unlike MP. Nonetheless, all three methods strongly support a group within the AC composed of Delgadobius, Inesius, Rhaegalius, Konradus, and Yuracarus (BS = 4, SH-aLRT = 94.7/UFB = 95, PP = 0.99). The monophyly of this group is also supported by one exclusive (30.1, male protarsomeres 2 and 3 sub-bilobed) and four non-exclusive (8.1*, head with protruding eyes; 31.2, male protarsomeres 2 and 3 subtriangular, i.e., flattened and widened distally; 32.1, male protarsomeres 2 and 3 with discal setae on ventral surface; 51.0, abdominal terga 3–7 almost glabrous) synapomorphies. Within this group, both MP and ML resolve Konradus as the sister group to the remaining taxa which appear together in a clade only weakly supported by MP (BS =1) as well as one exclusive (36.0, protarsomeres 2–3 without marginal setae on ventral surface) and two non-exclusive (29.1, prosternum with longitudinal ridge more or less rounded, from obtuse ridge to smooth longitudinal prominence of prosternum; 63.1, median lobe of aedeagus with apical fourth gradually narrowed in dorsal view) synapomorphies. Within this clade, Delgadobius appears as the sister-group to a subclade composed of Inesius, Rhaegalius and Yuracarus, weakly supported by MP and ML (BS = 4, SH-aLRT = 82.2/UFB = 91) as well as one exclusive (42.1*, mesotarsomere 2 with discal setae on ventral surface) and two non-exclusive (1.1*, body with metallic coloration limited to head, thorax and elytra; 23.1*, pronotum with lateral margins constricted before reaching posterior angles) synapomorphies. Within this subclade, the relationship between Inesius + Rhaegalius is strongly supported as a monophyletic group by all methods (BS = 5, SH-aLRT = 93/UFB = 97, PP = 1) alongside three exclusive (31.0, male protarsomeres 2 and 3 rectangular, i.e., flattened and rather transverse; 38.1, profemur sexually dimorphic, i.e., wider in males; 67.1, female genitalia with accessory sclerite linear) and three non-exclusive (60.0, paramere fused to median lobe only at base and 61.2, plate-like; 64.0, aedeagus with median lobe acute apically in dorsal view) synapomorphies, with Yuracarus as the sister group. Both Konradus and Yuracarus appear as monophyletic groups, strongly supported by all three methods (Konradus: BS = 3, SH-aLRT = 91.6/UFB = 97, PP = 0.97; Yuracarus: BS = 10?, SH-aLRT = 97.7/UFB = 98, PP = 1). Konradus is also defined by two exclusive (1.2*, entire body with metallic coloration; 11.0*, head with posterior angles almost obsolete) and one non-exclusive (43.0, elytra slightly punctate). Yuracarus is defined by three exclusive (39.1*, male profemora distinctly angulate medially; 41.0, mesoventrite with intercoxal ridge straight; 53.1*, abdominal tergum 8 with hind margin angulate medially in both sexes) and four non-exclusive (9.1, maximum width of head at its halfway between the anterior and posterior margin; 25.1, pronotum with posterior angles distinctly obtuse; 27.0, pronotum with large lateral setiferous puncture situated very close to superior marginal line of pronotum or at a distance no more than three times its diameter; 43.0*, elytra slightly punctate) synapomorphies.
Internal relationships within Konradus are weakly supported only by MP, with K. actinus as the sister taxon to the remaining three species (BS = 1) within which K. cuscensis and K. trescrucensis are sister taxa. In contrast, internal relationships within Yuracarus are supported by all three methods, with exception of K. diamantinus + K. yunguensis, unsupported by ML. Within Yuracarus, Y. cosnipatensis appears as the sister group to the remaining three species (BS = 3, SH-aLRT = 90.5/UFB = 97, PP = 0.85).
For the most relevant nodes (see Fig.
4. Discussion
The revision of type and non-type material collected from the Andes of Ecuador, Peru, and Bolivia, and deposited in entomological collections, led to an increase in the number of species in the previously monotypic and recently described genus Konradus, as well as the discovery of a new genus, Yuracarus. Additionally, two species originally described as Philonthus (Ph. actinus and Ph. diamantinus), but previously identified as “false” Philonthus species (e.g.,
Furthermore, within the subtriangular protarsomere clade, two additional traits emerge as particularly significant when unambiguously optimized. These include the presence of an accessory sclerite located along and between the gonocoxites and tergum 10, found in all genera of this group, including Konradus, and the dense, distinctly long discal setae of the protarsomeres 2 and 3, present in all genera except Konradus. While these traits, particularly the discal setae, may show dubious homology when observed outside this group, their uniqueness within the NL and the AC supports their inclusion as key synapomorphies. It should be noted that these derived characters were also observed in species outside the AC, within Philonthina. For instance, the occurrence of discal setae (which are also long and dense in some species of Cafius Stephens, 1829) and the accessory sclerite (which appears in Ph. splendens and some species of Paederomimus). However, the non-homology of the discal setae between the Holarctic representatives of Philonthina (including Cafius) and members of the AC has been previously proposed (
A possible hypothesis to explain the presence of dense, distinctly long discal setae on the ventral surface of the protarsomeres in Philonthina species inhabiting coastal (e.g., Cafius) or cloud forest (e.g., Yuracarus, Inesius) environments is that these structures may serve as an adaptation to enhance grip and stability on slippery, dynamic substrates, such as wet rocks, dense vegetation, or seaweed. Long setae may help prevent displacement by precipitation or water currents and facilitate the sensory detection of prey movements in fluctuating environments. In contrast, species of Philonthus living in more stable habitats, such as compost or forest soils, exhibit shorter setae, which are sufficient for maintaining traction and detecting vibrations in less dynamic conditions. This contrast suggests that setae length may be functionally linked to the stability and moisture characteristics of the beetle’s habitat. Thus, if a trait that might initially appear convergent (species of Cafius versus those of Yuracarus and allied genera) is present in the common ancestor of a group and accompanies the radiation of that group (i.e., Delgadobius, Inesius, Rhaegalius, and Yuracarus), it would no longer be regarded as convergence. Building on this, the dense, long discal setae present in the protarsomeres of Cafius and the aforementioned group are not homologous between these two groups, as they likely evolved independently in response to similar ecological pressures, particularly those associated with dynamic and slippery substrates such as coastal rocks, seaweeds, or the often wet canopy vegetation typical of cloud forests. However, within the “setose protarsomere” clade the presence of these setae would be homologous, as they are inherited from their common ancestor and provide consistent phylogenetic signal. This further supports the hypothesis that these characters define a distinct and well-supported clade within the AC of the NL.
In conclusion, the presence of these setae, along with other derived characters, can be used to recognize and define a specific clade within the AC, strengthening the case for their phylogenetic relevance and aiding in the systematic classification of these taxa. Future studies should continue to explore the evolutionary and ecological significance of these traits, further elucidating the mechanisms driving diversification within this fascinating group of beetles in the Neotropical region.
5. Funding
This study was financed by Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) [PIP DD87 (M. Chani-Posse and F. Agrain)] and Agencia Nacional de Promoción Científica y Tecnológica (ANPCyT) – Fondo para la Investigación Científica y Tecnológica (FONCYT) [PICT 2019-03121 (M. Chani-Posse and F. Agrain)].
6. Competing interests
The authors have declared that no competing interests exist.
7. Acknowledgements
We gratefully acknowledge the help received from all museum curators here when loaning the specimens. We also thank the Willi Hennig Society for the free use of TNT. MRM thanks to Conservación Amazónica – ACCA for the scholarship awarded to carry out field trips at the Estación Biológica Wayqecha (2017 and 2019) where Yuracarus cosnipatensis was collected.
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Supplementary material
File S1
Data type: .nex
Explanation notes: Morphological matrix (Mesquite).