Research Article |
Corresponding author: Andrey V. Frolov ( afrolov@zin.ru ) Academic editor: Patrice Bouchard
© 2023 Andrey V. Frolov, Lilia A. Akhmetova, Jhon César Neita-Moreno.
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.
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://doi.org/10.3897/zookeys.1166.102813
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In the Neotropics, orphnine scarab beetles are represented by the endemic tribe Aegidiini Paulian, 1984 with five genera and over 50 species. Phylogenetic analysis based on morphological characters of all supraspecific taxa of Orphninae showed that Aegidiini 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.
Caqueta moist forests, Colombia, new species, new subtribes, orphnines, Peru, Peruvian Yungas, phylogeny, scarabs
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 (
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 (
The material used in this work is housed in the collection of the Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, Russia (
The maximum parsimony (MP) analyses were conducted in TNT 1.6 (
Ingroup and outgroup
In the ingroup, we included all generic taxa of the Orphninae with the following 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 (
The character states are based on the previous phylogenetic analysis (
The characters 14, 17, 37, 41, 42 (
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
The parsimony phylogenetic analysis yielded six most parsimonious trees 83 steps long (Suppl. material
Phylogeny of Orphninae based on parsimony analysis of the revised character states from Frolov (
Family Scarabaeidae Latreille, 1802
Subfamily Orphninae Erichson, 1847
Aegidium.
Small to medium-sized beetles (body length 5–20 mm), brown to black colored without pattern, more or less densely punctate, smooth or densely setose. Mandibles subsymmetrical, without lateral processes, distinctly 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 bordering the excavation near anterior margin (lateral pronotal processes), and with a tubercle or small horn medially on the anterior margin (anterior pronotal process); these characters are subject to allometric variability and may not be developed in some males. Females have a convex pronotum without armature 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 elytra. 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 triangular, 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. Parameres symmetrical, relatively long, apices tapering or curved downwards, with or without setae; a few species have complex, feeble sclerotised processes 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.
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).
Endemic to South and Central America.
Aegidinus Arrow, 1904.
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, convex to depressed medially in females; anterior margin of pronotum in males with a tubercle or horn medially. Propleura with carinae separating anterolateral 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. Metepisternon 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 ventroapically in males. Stridulatory file with wide carinae medially becoming much narrower and denser proximally. Phallobase tube shaped with strongly sclerotised 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 variably shaped, sometimes angulate or with a process mediabasally; coxites variably shaped, with armature sort robust spinules in some species mediabasally, stili distinct, variably shaped, or indistinct, not separated from coxites.
Only type genus, Aegidinus Arrow, 1904 (16 spp).
Endemic to South America.
Aegidinus alexanderi sp. nov. is most similar to A. teamscaraborum 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.
Aegidinus alexanderi sp. nov. (A, D, E male, holotype B female, paratype) and A. teamscaraborum (G) A, B habitus 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.).
Holotype. Male at
Male, holotype (Fig.
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.
Pronotum with widely rounded lateral margins, narrower than elytra, 1.6 times wider than length. Posterior angles widely rounded. Anterior margin bordered, 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 anteromedially 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 basally 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 tarsomere. Femora almost impunctate.
Abdomen
ventrally irregularly punctate, pubescent, with sparse, long setae. Abdominal sternite 8 medially slightly longer than sternites 4–7 combined. Pygidium invisible from above, with slightly truncate apex in caudal view. Plectrum triangular with rounded apex, wider than long. Stridulatory file (Fig.
Aedeagus. Phallobase without ventroapical plate. Parameres short (about 0.4 length of phallobase), curved downwards (Fig.
Female (Fig.
The body length of the examined specimens varies from 7.8–8.5 (males) and from 7.5–9.0 (females). Head and pronotal armature in one male paratype poorly developed with a small frontoclypeal tubercle and shallow pronotal fossa medially.
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.
The new species is named after Alexander Petrov (Moscow) who collected and kindly donated us the specimens.
Aegidinus elbae sp. nov. is similar to A. colbyae
Holotype. Male at
Male (Fig.
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 bordered, 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 anteromedially 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 basally 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 tarsomere. Femora almost impunctate.
Aedeagus. Phallobase without ventroapical plate. Parameres long (about 0.7 length of phallobase). Parameres symmetrical, of complex shape (Fig.
Female (Fig.
The species is known from a single locality in Caquetá, Colombian Amazonia (Fig.
The species is dedicated to Lic. Elba Moreno de Neita, mother of JCNM, to honor her memory.
1 | Parameres separated into dorsomedial and ventrolateral lobes | 2 |
– | Parameres not separated into dorsomedial and ventrolateral lobes | Aegidinus cornutus Colby, 2009 |
2 | Phallobase with ventroapical plate | 3 |
– | Phallobase without ventroapical plate | 6 |
3 | Parameres symmetrical | 4 |
– | Parameres asymmetrical | 5 |
4 | Ventrolateral lobe of paramere with subapical tooth | Aegidinus howdenorum Colby, 2009 |
– | Ventrolateral lobe of paramere without subapical tooth | Aegidinus guianensis (Westwood, 1845) |
5 | Parameres longer, more asymmetrical; ventroapical plate of phallobase longer than wide; protibia without medioapical tooth | Aegidinus noriegai Frolov, Akhmetova & Vaz-de-Mello, 2019 |
– | Parameres shorter, less asymmetrical; ventroapical plate of phallobase wider than long; protibia with medioapical tooth | Aegidinus candezei (Preudhomme de Borre, 1886) |
6 | Mediobasal margins of dorsomedial lobes of parameres feebly sclerotized, membranous; protibia with medioapical tooth | 7 |
– | Mediobasal margins of dorsomedial lobes of parameres strongly sclerotized; protibia without medioapical tooth | 10 |
7 | Ventrolateral lobes of parameres long and slender (in lateral view), reasonably longer than dorsomedial lobes | Aegidinus steinheili (Harold, 1880) |
– | Ventrolateral lobes of parameres triangular and obtuse in lateral view, not longer than dorsomedial lobes | 8 |
8 | Ventrolateral lobes of parameres as long as dorsomedial lobes | Aegidinus petrovi Colby, 2009 |
– | Ventrolateral lobes of parameres reasonably shorter than dorsomedial lobes | 9 |
9 | Parameres with proximal and distal lobes more separated and proximal lobes shorter in lateral view | Aegidinus teamscaraborum Colby, 2009 |
– | Parameres with proximal and distal lobes less separated and proximal lobes longer in lateral view (Fig. |
Aegidinus alexanderi Frolov, Akhmetova & Neita-Moreno, sp. nov. |
10 | Dorsal sides of parameres strongly overlapping and separated by slit | Aegidinus simulates Colby, 2009 |
– | Dorsal sides of parameres less overlapping and not separated by slit | 11 |
11 | In lateral view, parameres longer (about 0.7 length of phallobase) and abruptly separated into apical and basal parts (Fig. |
Aegidinus elbae Neita-Moreno, Akhmetova & Frolov, sp. nov. |
– | In lateral view, parameres shorter (about 0.4 length of phallobase), not abruptly separated into apical and basal parts | 12 |
12 | Dorsal processes of parameres carina-shaped | Aegidinus colbyae Frolov, Akhmetova & Vaz-de-Mello, 2019 |
– | Dorsal processes of parameres tooth or spur-shaped | 13 |
13 | Dorsal processes of parameres long, spur-shaped | Aegidinus brasiliensis Arrow, 1904 |
– | Dorsal processes of parameres short, tooth-shaped | Aegidinus howeae Colby, 2009 |
Although the monophyly of Aegidiini was not questioned (
The analysis reported here, based on the expanded and verified set of characters and all nominal supraspecific Orphninae taxa, provides strong support for the two lineages of the Aegidiini. One lineage includes Aegidium and three related genera. It has a unique synapomorphy, the hole connecting mesocoxal cavities, and a non-unique synapomorphy, the absence of the transverse keel on hind tibiae, shared with Stenosternus. However, the latter state could be gained independently by the New World and Old World taxa. The second lineage included the genus Aegidinus and is characterised by two unique synapomorphies, mandibles with long processes on the outer sides and two procoxal hollows.
To make the classification better reflect the phylogenetic relations of the taxa in questions, and specifically to distinguish the Aegidium group lineage as a separate taxon we introduced a subtribal system for Aegidiini. The nominotypical subtribe, Aegidiina, includes Aegidium, Paraegidium, Aegidiellus and Onorius, and a new monotypical subtribe, Aegidinina subtr. nov. is erected to accommodate Aegidinus.
Aegidinus now comprises 15 species and is the second most species-rich genus of the South American Orphninae, after Aegidium. The bionomy of its species is virtually unknown and almost all species were recorded from a small series of specimens. It is possible that more species are yet to be described, specifically from the Andes, which is apparently the centre of diversity and diversification of the genus. Our results also suggest that the Aegidinus is composed of a few lineages, which may necessitate establishing subgeneric or species group classification, however the analysis of the phylogenetic relations of the taxa at species level is outside the scope of the present contribution.
We are thankful to Alexander Petrov (Moscow) who provided the specimens of the new Aegidinus species. We are especially thankful to Andrew Smith and Mario Cupello for the comments that considerably improved the draft manuscript. The study was performed in the framework of the Russian State research project 122031100272-3.
No conflict of interest was declared.
No ethical statement was reported.
No funding was reported.
Conceptualization: AF, investigation: all authors; data curation: all authors; writing original draft: all authors; review and editing draft: all authors; project administration: AF.
Andrey V. Frolov https://orcid.org/0000-0002-6724-6828
Lilia A. Akhmetova https://orcid.org/0000-0002-2151-1278
Jhon César Neita-Moreno https://orcid.org/0000-0003-2998-2063
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Character states
Data type: morphological (word document)
Explanation note: Morphological character states for phylogenetic analysis.
Character matrix
Data type: phylogenetic (word document)
Phylogenetic trees
Data type: images (PDF file)
Explanation note: Six most parsimonious trees, resulted from phylogenetic analysis.