#ZooKeys ZooKeys 1166: 33-47 (2023) DOI: 10.3897/zookeys.1166.102813 Research Article Phylogenetic analysis of the Neotropical scarab beetle tribe Aegidiini (Coleoptera, Scarabaeidae, Orphninae) with description of new taxa Andrey V. Frolov™®, Lilia A. Akhmetova™®, Jhon César Neita-Moreno2® 1 Laboratory of Insect Systematics, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St. Petersburg 199034, Russia 2 Instituto de Investigaciones de Recursos Bioldgicos Alexander von Humboldt, Claustro de San Agustin, Boyacda, Colombia Corresponding author: Andrey V. Frolov (afrolov@zin.ru) OPEN Qaccess Academic editor: Patrice Bouchard Received: 2 March 2023 Accepted: 15 May 2023 Published: 6 June 2023 ZooBank:https://zoobank.org/94B60F4A- 8430-4FDA-9C92-1B9C66833E83 Citation: Frolov AV, Akhmetova LA, Neita-Moreno JC (2023) Phylogenetic analysis of the Neotropical scarab beetle tribe Aegidiini (Coleoptera, Scarabaeidae, Orphninae) with description of new taxa. ZooKeys 1166: 33-47. https://dol.org/10.3897/ zookeys.1166.102813 Copyright: © Andrey V. Frolov et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract In the Neotropics, orphnine scarab beetles are represented by the endemic tribe Aegi- diini Paulian, 1984 with five genera and over 50 species. Phylogenetic analysis based on morphological characters of all supraspecific taxa of Orphninae showed that Ae- gidiini is comprised of two lineages. New subtribes, Aegidiina subtr. nov. (Aegidium Westwood, 1845, Paraegidium Vulcano et al., 1966, Aegidiellus Paulian, 1984, and Onorius Frolov & Vaz-de-Mello, 2015, and Aegidinina subtr. nov. (Aegidinus Arrow, 1904) are proposed to better reflect this phylogeny. Two new species of Aegidinus are described: A. alexanderi sp. nov., from the Yungas in Peru and A. elbae sp. nov. from the Caqueta moist forests ecoregion in Colombia. A diagnostic key to Aegidinus species is given. Key words: Caqueta moist forests, Colombia, new species, new subtribes, orphnines, Peru, Peruvian Yungas, phylogeny, scarabs Introduction The scarab beetles of the subfamily Orphninae are distributed mostly in the tropics of the southern continents. In the Neotropics, they are represented by the endemic tribe Aegidiini Paulian and comprise five genera and over 50 species (Paulian 1984; Colby 2009; Frolov and Vaz-de-Mello 2015; Frolov et al. 2017a, b, c; Rojkoff and Frolov 2017; Frolov et al. 2019). In the previous phylogenetic analysis of the Orphninae (Frolov 2012), some characters were misinterpreted due to the limited material then available. After this prelimi- nary analysis was published, a new genus of the South American Orphninae was also described (Frolov and Vaz-de-Mello 2015). The aim of the present work, apart from the description of a new species of Aegidinus Arrow, is to provide the results of the phylogenetic analysis of the tribe Aegidiini based on a verified and expanded set of morphological characters of all nominal supraspecific taxa of the Aegidiini and make the classification better reflect the phylogenetic relations of the taxa in question by introducing a subtribal level with two new taxa. 33 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini The genus Aegidinus currently comprises 14 species distributed in South America, mostly in the Amazon and Guiana moist forest regions to the Yungas in the west, and in Trinidad Island (Colby 2009; Frolov et al. 2019). Recently we had an opportunity to examine two series of Aegidinus specimens from south-eastern Peru. The series included both males and females and are sim- ilar to A. teamscaraborum Colby, 2009, yet the males have a different shape of the parameres. Below the new species is described. Material and method The material used in this work is housed in the collection of the Zoological In- stitute, Russian Academy of Sciences, Saint-Petersburg, Russia (ZIN), Instituto Alexander von Humboldt, Villa de Leyva, Boyaca, Colombia (IAvH), and Cana- dian Museum of Nature, Ottawa, Canada (CMN). Morphological terminology follows Frolov (2012) and Frolov et al. (2016). In the new species descriptions, labels of the type specimens are cited verbatim and separated by a slash and our comments are in square brackets. Preparation of specimens, digital imag- es and locality maps follow Akhmetova and Frolov (2022). The maximum parsimony (MP) analyses were conducted in TNT 1.6 (Golo- boff and Morales 2023) using the “traditional search” option to find the most parsimonious trees (MPTs). The following parameters were used: memory set to hold 1 000 000 trees; tree bisection—reconnection (TBR) branch-swap- ping algorithm with 1000 replications saving ten trees per replicate; ze- ro-length branches collapsed after the search. All character states were treated as unordered and equally weighted. Autapomorphic characters were deactivated before the parsimony analysis. Bremer support was calculated using the TNT Bremer function, using suboptimal trees up to 20 steps lon- ger. For character mapping, Winclada v.1.00.08 (Nixon 2002) was used with unambiguous optimization. Results Phylogenetic analysis of the Aegidiini Ingroup and outgroup In the ingroup, we included all generic taxa of the Orphninae with the fol- lowing exceptions. The monotypic genera Hybaloides Quedenfeldt, 1884 and Craniorphnus Kolbe, 1895, known from single type specimens, are not included since they are based on misidentified Orphnus species (unpublished data of the authors). The two genera, Onorius Frolov & Vaz-de-Mello, recently described from the Andes (Frolov and Vaz-de-Mello 2015), and the central African Cerhomalus Quedenfeldt, 1884, restored as a genus distinct from Orphnus (Frolov and Akhmetova 2021), are added to the list of ingroup taxa used by Frolov (2012). From the outgroup, we excluded the distantly related taxa of the family Hybosoridae and included the two species of the genus Allidiostoma Arrow, 1904. The latter is a member of the small, olygotypic sub- family Allidiostomatinae Arrow, 1904, distributed in southern South America ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 34 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini (Ocampo and Colby 2009). There is evidence that Allidiostomatinae might be a sister group of the Orphninae (Ocampo and Hawks 2006; Ocampo et al. 2010; Neita-Moreno et al. 2019). Character states and their codes The character states are based on the previous phylogenetic analysis (Frolov 2012) with the following modifications: Characters excluded The characters 14, 17, 37, 41, 42 (Frolov 2012) were excluded because they be- come uninformative after Hybosoridae was excluded from the analysis. Char- acters 24 and 25 are excluded since they are related to flightlessness which occurs in many Orphninae taxa and apparently evolved many times in different lineages (Frolov and Akhmetova 2020). Character 31 is excluded because it is a character of sexual dimorphism found in different non-related taxa (i.e., Scarabaeinae dung beetles of the genera Macroderes Westwood, 1842 and Xinidium Harold, 1869). Characters added Parameres: symmetrical (0), asymmetrical (1). Stridulatory ridges: straight (0), distinctly curved posteriad (1). Phallobase protruding ventroapical plate: absent (0), present (1). Mediobasal margins of parameres: feebly sclerotised (0), strongly sclerotised (1), strongly sclerotised and serrate (2). Mandibles visible from above: yes (0), no or feebly (1). Labrum visible from above: yes (0), no (1). Tarsi: slender (0), robust (1). Paramere apices: glabrous (0), with short setation (1), with long setation (2). Tubercle on anterior margin of pronotum in female: absent (0), present (1). Clypeus anteriorly in males: not bilobate or bifurcated (0), bilobate or bifurcated (1). Dorsum of body: minutely setose or glabrous (0), densely pubescent (1). Elytron, longitudinal keels: no (0), 2 (1), 1 (2). Phallobase: membranous ventro-proximally (0), tube shaped (1). Phallobase ventrally: entirely membranous (0), sclerotised apically (1). Phallobase, ventroapical sclerotization: 1 large sclerite (0), 2 swollen sclerites (1). The complete list of the character states and the matrix are provided in the Suppl. materials 1, 2. Tree topologies The parsimony phylogenetic analysis yielded six most parsimonious trees 83 steps long (Suppl. material 3). The trees show a similar topology differing most- ly in the position of some Old World lineages. The strict consensus is shown in the Fig. 1. ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 35 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini Allidiostoma ramosae Allidiostoma sp. 17 31 , 0 Orphnus giganteus 3 11 15 Goniorphnus felschei 50 Pp Cerhomalus 9 Orphnus macleayi 10 16 wed Orphnus sansibaricus Pseudorphnus hyboni = 10 20 _ on Triodontus nitidus = 4 18 10 23 oO. 0 Renorphnus clementi co 8 9 10 16 cy 6 10 21 22 24 25 29 30 32 35 36 O Madecorphnus falcatus 19 27 28 40 44 48 50 51 = = Stenosternus costatus 2 12 25 44 50 CL J CL) 4 5 10 15 17 D Hybalus digitalis [_ CJ 16 Chaetonyx robustus 13 44 45 Aegidiellus alatus Aegidiellus zezaoi ine) 19 46 47 Paraegidium costalimai = 28 34 Paraegidium barretoi = 7 =) Pee eB) N41 42 43 Onorius inexpectatus < Onorius sp. Ag Aegidium columbianum Aegidium howdeni 9 38 49 Aegidinus cornutus Aegidinus petrovi Aegidinus teamscaraborum Aegidinus steinheli B26 Aegidinus alexanderi Aegidinus simulates Aegidinus colbyae Aegidinus howeae Aegidinus guianensis 37 Aegidinus candezei Aegidinus noriegai Figure 1. Phylogeny of Orphninae based on parsimony analysis of the revised character states from Frolov (Frolov 2012) with additional taxa. The strict consensus of six most parsimonious trees (83 steps, ci 84, ri 92). Black boxes — unique synapomorphies, white boxes — nonexclusive synapomorphies (homoplasies). Taxonomical accounts Family Scarabaeidae Latreille, 1802 Subfamily Orphninae Erichson, 1847 Tribe Aegidiini Paulian, 1984 Subtribe Aegidiina Paulian, 1984 Type genus. Aegidium. Diagnosis. Small to medium-sized beetles (body length 5-20 mm), brown to black colored without pattern, more or less densely punctate, smooth or ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 36 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini densely setose. Mandibles subsymmetrical, without lateral processes, dis- tinctly or feebly protruding past anterior margin of frontoclypeus in dorsal view. Labrum exposed or hidden under clypeus in dorsal view. Frontoclypeus symmetrical or subsymmetrical, without tubercles, horns or ridges, or in males with variably shaped bilobate anterior frontoclypeal process. Pronotum of males may be with deep excavation in the middle, with 2 horns or ridges bor- dering the excavation near anterior margin (lateral pronotal processes), and with a tubercle or small horn medially on the anterior margin (anterior prono- tal process); these characters are subject to allometric variability and may not be developed in some males. Females have a convex pronotum without arma- ture or pronotum impressed anteriorly on disc and with a tubercle medially on anterior margin. Propleurae with carinae separating anterolateral areas from basal area. Scutellum narrowly rounded apically, about 1/8-1/13 length of el- ytra. Elytra convex, with marked humeral umbones (except for brachypterous species). Surface flat or with two low ridges in basal half; the ridges may be more or less convex, smooth, to almost indistinct. Pubescence of dorsal side indistinct or dense. Wings fully developed or vestigial. Metepisternon triangu- lar, its posterior angle rounded to triangular and situated in distinct concavity of epipleuron. Mesocoxal cavities connected by a hole. Protibiae with three outer teeth, somewhat serrate basad of the teeth, with a smaller, medial tooth in majority of males. In males, anterior spur is absent. Each procoxae with one elongate hollow. Mesotibiae with or without a tuft of setae ventroapically in males. Stridulatory file with relatively fine, evenly spaced carinae. Phallobase tube shaped with strongly sclerotized ventral side but without differentiation of ventral and dorsal sclerites; ventroapical plate absent or present. Param- eres symmetrical, relatively long, apices tapering or curved downwards, with or without setae; a few species have complex, feeble sclerotised process- es on the parameres lateroapically. Endophallus without armature or with a small group of spinules; in one species of Aegidium there is a sclerite with two large curved spines. Spiculum gastrale T-, Y- or V-shaped, with setae on apical plate. Subcoxites oval, with dense, long setae mediabasally; coxites triangular, long, with dense short setae mediabasally and sparse long setae apically; stili distinct, elongated, or not separated from coxites. Taxon composition. The subtribe is comprised of Aegidium Westwood, 1845 (25 spp), Paraegidium Vulcano et al., 1966 (6 spp), Aegidiellus Paulian, 1984 (3 spp) and Onorius Frolov & Vaz-de-Mello, 2015 (2 spp). Distribution. Endemic to South and Central America. Aegidinina Frolov, AAhmetova & Neita-Moreno, subtr. nov. https://Zoobank.org/CE492F83-9066-421E-B89B-53A36C1B1C4B Type genus. Aegidinus Arrow, 1904. Diagnosis. Body small to mid-sized (length 6 to 12 mm), reddish brown to dark brown. Mandibles subsymmetrical, with long processes on the outer sides. Clypeus with tubercle or horn on anterior margin medially in males, without horn in females. Pronotum variably excavated medially in males, con- vex to depressed medially in females; anterior margin of pronotum in males with a tubercle or horn medially. Propleura with carinae separating antero- ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 37 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini lateral areas from basal area. Scutellum narrowly rounded posteriorly, about 1/12 length of elytra. Elytra convex, with marked humeral umbones and striae marked with elongated punctures, surface smooth. Wings fully developed. Me- tepisternon triangular, its posterior angle rounded to triangular and situated in distinct concavity of epipleuron. Mesocoxal cavities not connected by a hole. Protibiae with three outer teeth, somewhat serrate basad of the teeth, with a smaller, medial tooth in majority of males. In males, anterior spur is absent. Each procoxa with two hollows. Mesotibiae without a tuft of setae ventroap- ically in males. Stridulatory file with wide carinae medially becoming much narrower and denser proximally. Phallobase tube shaped with strongly scle- rotised ventral side but without differentiation of ventral and dorsal sclerites; ventroapical plate absent. Parameres relatively short, with complex shape but without feeble sclerotised processes, apices without setae; in some species parameres strongly asymmetric. Endophallus with relatively well-developed armature consisting of a few groups of spinules, sometimes of different size. Spiculum gastrale Y-shaped, without setae on apical plate. Subcoxites vari- ably shaped, sometimes angulate or with a process mediabasally; coxites variably shaped, with armature sort robust spinules in some species media- basally, stili distinct, variably shaped, or indistinct, not separated from coxites. Taxon composition. Only type genus, Aegidinus Arrow, 1904 (16 spp). Distribution. Endemic to South America. Aegidinus alexanderi Frolov, Akhmetova & Neita-Moreno, sp. nov. https://zoobank.org/710FOE2D-0F47-4CB6-8355-2A71 734E930E Fig. 2A—H Differential diagnosis. Aegidinus alexanderi sp. nov. is most similar to A. teams- caraborum Colby, 2009, but differs from it in the shape of the parameres having proximal and distal lobes less separated and proximal lobes longer in lateral view (Fig. 2E—-G). It should be noted that the differences between the two species are smaller than between most other Aegidinus species, implying their close relation- ships. It is possible that a thorough sampling in the Yungas will provide clear ev- idence of their allopatric or parapatric distribution and examination of molecular markers will show that the genetic distance between them is characteristic for sub- species rather than species; in this case, their status may be changed. Until such data are available, we suggest that these taxa are considered distinct species. Type material. Holotype. Male at ZIN labeled “JUNIN: Satipo Prov., 5 km NNE Puerto Ocopa, left bank of Perené River, near Canan Eden village, 1100 ma.s.l., vill. 8.111.2008. A.Petrov leg [FIDE6071]”. Paratypes. One male and one female [FIDE6072, FIDE6073] at CMN and two females [FIDE6074, FIDE6075] at ZIN with the same data as the holotype; one male and three females at ZIN labeled “PERU: Junin, 16 km NW Satipo, rio Venado, 1150 m 11°11.677'S, 74°46.137'W 13.11.2010 A. Petrov leg. [FIDE6076—-FIDE6079]”. Description. Male, holotype (Fig. 2A, D, E). Body length 8.4 mm. Colour uniformly dark brown. Frontoclypeus wide, with convex anterior margin, slightly angulate laterally, somewhat crenulate. Genae small, slightly protruding past eyes. Frontal suture indistinct. Frontoclypeus with short conical horn rounded apically. ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 38 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini 1mm () Aegidinus alexanderi sp. nov. {|_| Aegidinus teamscaraborum H F G Figure 2. Aegidinus alexanderi sp. nov. (A, D, E male, holotype B female, paratype) and A. teamscaraborum (G) A, B hab- itus C stridulatory file, SEM D parameres in dorsal view E aedeagus in lateral view F, G paramere outline in lateral view (not to scale) H distributional record map (red symbols indicate holotype localities, gray squares indicate localities of A. teamscaraborum paratypes, which may belong to A. alexanderi sp. nov.). See eee eee eee Pronotum with widely rounded lateral margins, narrower than elytra, 1.6 times wider than length. Posterior angles widely rounded. Anterior margin bor- dered, border interrupted medially, with feeble gibbosity. Base of pronotum not bordered, with a few large rounded punctures laterally and a few small medially. Pronotal disc feebly excavated anteromedially, with two gibbosities in center. Pronotum punctate with a few large rounded punctures laterally and anterome- dially and with minute, feebly visible punctures throughout. Scutellum subtriangular, narrowly rounded posteriorly, about 1/11 length of elytra. Elytra almost as long as wide, widest medially and rounded apically, with humeral and apical humps. First elytral stria as continuous line, connected ba- ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 39 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini sally with undulate line from scutellum to humeral hump. Other striae marked with rows of sparse punctures; punctures somewhat V- and comma-shaped on basal part of elytra, becoming smaller towards apices. Macropterous. Legs. Protibiae with 3 outer teeth, without medioapical tooth. Lateral margin basad of outer teeth not crenulate. Apical spur of protibia absent. Middle and hind legs similar in shape; metafemora and metatibiae about 1/8 longer than the mesofemora and mesotibiae. Mesotibia and metatibiae with 2 apical spurs, inner margin almost straight, outer margin with 1 transverse keel. Upper spur of hind tibiae as long as two basal tarsomeres. Claws 1/3 length of apical tar- somere. Femora almost impunctate. Abdomen ventrally irregularly punctate, pubescent, with sparse, long setae. Abdominal sternite 8 medially slightly longer than sternites 4-7 combined. Py- gidium invisible from above, with slightly truncate apex in caudal view. Plec- trum triangular with rounded apex, wider than long. Stridulatory file (Fig. 2C) with wide carinae medially becoming much narrower and denser proximally. Aedeagus. Phallobase without ventroapical plate. Parameres short (about 0.4 length of phallobase), curved downwards (Fig. 2D, E). Parameres with prox- imal lobes reasonably longer than sinuation between proximal and distal lobes in lateral view (Fig. 2E). Endophallus with 3 groups of spinules. Female (Fig. 2B) differs from the male in having a relatively smaller prono- tum without armature, frontoclypeus without process, and short but distinct protibial spur. Paratypes and variability. The body length of the examined specimens var- ies from 7.8—-8.5 (males) and from 7.5—9.0 (females). Head and pronotal arma- ture in one male paratype poorly developed with a small frontoclypeal tubercle and shallow pronotal fossa medially. Distribution. This species is known from two localities in Satipo Province in central Peru, mostly within the Peruvian Yungas ecoregion and on the border with Southwest Amazon moist forests ecoregion (Fig. 2H). The records of the para- types A. teamscaraborum from the Peruvian Yungas (Fig. 2H, gray squares) are based on females only therefore may belong to A. alexanderi sp. nov. The holo- type and other paratypes of A. teamscaraborum originate from Bolivian Yungas. Etymology. The new species is named after Alexander Petrov (Moscow) who collected and kindly donated us the specimens. Aegidinus elbae Neita-Moreno, Akhmetova & Frolov, sp. nov. https://zoobank.org/89396271-30BC-4CA1-BC43-010C3871E8C3 Figa3ATE Differential diagnosis. Aegidinus elbae sp. nov. is similar to A. co/byae Frolov et al., 2019, A. brasiliensis Arrow, 1904 and A. howeae Colby, 2009 in having mediobasal margins of dorsomedial lobes of parameres strongly sclerotized, protibia without medioapical tooth, and dorsal sides of parameres less over- lapping, but differs from them in the parameres being distinctly longer and, in lateral view, abruptly separated into apical and basal parts (Fig. 3C). Type material. Holotype. Male at |AvH (Fig. 3A): “COLOMBIA, Caqueta, Sola- no PNN/ Chiribiquete, Rio Sararamano/Bosque Verde militar 300 m/0°14'47'N, ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 40 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini 1mm 0.5mm oho Aegidinus elbae sp. nov. a Aegidinus noriegai Bogota |_| Aegidinus steinheili tees ® & Aegidinus candezei A llayicencio A Aegidinus simulatus ai e Wscrak Guenea’ 7 Figure 3. Aegidinus elbae Neita-Moreno, Akhmetova & Frolov sp. nov. (A, C, D male, holotype B female, paratype) A, B hab- itus C aedeagus in lateral view D parameres in dorsal view E distributional record map. ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 4 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini 72°37'24'W pitfall T2/T1 3-5.iv.2000, E. Gonzalez Leg.”, “IAVH-E-256378”. Paratype. Female at I|AvH (Fig. 3B) with following data: “COLOMBIA, Caqueta, So- lano PNN/ Chiribiquete, Rio Sararamano/Bosque Verde militar 300 m/0°14'47'N, 72°37'24'W pitfall /3-5.iv.2000, E. Gonzalez Leg.”, “IAVH-E-256379”. Description. Male (Fig. 3A, C, D). Body length 9.6 mm. Colour uniformly dark brown. Anterior margin of frontoclypeus with horn rounded apically. Pronotum with widely rounded lateral margins, narrower than elytra, 1.6 times wider than length. Posterior angles widely rounded. Anterior margin bor- dered, border complete medially, with feeble gibbosity. Base of pronotum not bordered, with a few large rounded punctures laterally and a few small medially. Pronotal disc feebly excavated anteromedially, with two gibbosities in centre. Pronotum punctate with a few large rounded punctures laterally and anterome- dially and with minute, feebly visible punctures throughout. Scutellum subtriangular, narrowly rounded posteriorly, about 1/11 length of elytra. Elytra almost as long as wide, widest medially and rounded apically, with humeral and apical humps. First elytral stria as continuous line, connected ba- sally with undulate line from scutellum to humeral hump. Other striae marked with rows of sparse punctures; punctures somewhat V- and comma-shaped on basal part of elytra, becoming smaller towards apices. Macropterous. Legs. Protibiae with 3 outer teeth, without medioapical tooth. Lateral margin basad of outer teeth not crenulate. Apical spur of protibia absent. Middle and hind legs similar in shape; metafemora and metatibiae about 1/8 longer than the mesofemora and mesotibiae. Mesotibia and metatibiae with 2 apical spurs, inner margin almost straight, outer margin with 1 transverse keel. Upper spur of hind tibiae as long as two basal tarsomeres. Claws 1/3 length of apical tar- somere. Femora almost impunctate. Aedeagus. Phallobase without ventroapical plate. Parameres long (about 0.7 length of phallobase). Parameres symmetrical, of complex shape (Fig. 3C, D): in the lateral view, they are abruptly separated into apical and basal parts; basal parts with 2 acute processes, apical parts somewhat dilating, giving the shape of a cup, acute and curved at very apex. Female (Fig. 3B) differs from male in having relatively wider elytra, prono- tum and head without excavations and armature, and in having a protibial spur. Body length 8.7 mm. Distribution. The species is known from a single locality in Caqueta, Colom- bian Amazonia (Fig. 3E). Etymology. The species is dedicated to Lic. Elba Moreno de Neita, mother of JCNM, to honor her memory. Key to the species of Aegidinus Arrow (males) 1 Parameres separated into dorsomedial and ventrolateral lobes. ............... 2 — Parameres not separated into dorsomedial and ventrolateral lobes........... dilated dated scalunl acgedudeawissit ee NMG ua eea led Aegidinus cornutus Colby, 2009 2 Phallobase with ventroapical plate ................ccesccssssceesesrsecesseeecssseeeeessneeees 3 —- Phallobase without ventroapical plate................ ccc ecessscecssseeessseeeesssseeesens 6 ZooKeys 1166: 33-47 (2023), DOI: 10.3897/zookeys.1166.102813 42 Andrey V. Frolov et al.: Phylogenetic analysis of Aegidiini 10 17 12 13 RarAanenre SeS¥MEMC UAC All crt rece.ceoch.,nesevesecmmsneAmuaten ated ent sarees neneosercterredimcaws cael 4 PararnereS aS VM nme meal i... acectans dvencaeannslnteeeataalnaaantsieareneecenate date, 5 Ventrolateral lobe of paramere with subapical tooth..............cccceseeeseeeeees Me elie eatad weak etn eens Foie, co dite, Me aes eee Aegidinus howdenorum Colby, 2009 Ventrolateral lobe of paramere without subapical tooth .................ccceeees ieee ae start, before Peau eras, Aegidinus guianensis (Westwood, 1845) Parameres longer, more asymmetrical; ventroapical plate of phallobase longer than wide; protibia without medioapical tooth .............. cece eens boscteche dus ec 3 Aegidinus noriegai Frolov, AKhmetova & Vaz-de-Mello, 2019 Parameres shorter, less asymmetrical; ventroapical plate of phallobase wider than long; protibia with medioapical tooth ........... ee eeeeceesseeeeeneees Pe pire hnan Air Ehrich Aegidinus candezei (Preudhomme de Borre, 1886) Mediobasal margins of dorsomedial lobes of parameres feebly sclero- tized, membranous; protibia with medioapical tooth............... eee eens 7 Mediobasal margins of dorsomedial lobes of parameres strongly sclero- tized; protibia without Medioapical tOOth .......... ee eesscesessteeeseteeeeenneees 10 Ventrolateral lobes of parameres long and slender (in lateral view), reason- ably longer than dorsomedial lobes.....Aegidinus steinheili (Harold, 1880) Ventrolateral lobes of parameres triangular and obtuse in lateral view, not longershan; dorsomedial | GWCS AS ...