Vasily V. Grebennikov’s research while affiliated with Canadian Food Inspection Agency and other places

The tribe Raymondionymini has long been neglected in phylogenetic studies. The tribe is characterized by uncertain monophyly, fluctuating taxonomic status, and a composition prone to instability. All raymondionymine weevils are wingless and have eyes either completely absent or, rarely, consisting of a single ommatidium. With body lengths predominantly below three millimeters, they inhabit deep soil environments and are infrequently collected. The core of this tribe comprises nine genera distributed in Europe and around the Mediterranean region and encompassing 76 species, while six additional genera include 17 species distributed in USA (California), Mexico, Ecuador, Venezuela, Russian Far East, and Madagascar. Here, we present eight new mitogenomes, complemented by one publicly available, encompassing all but two Mediterranean genera of raymondionymine weevils. We used publicly available Curculionoidea mitogenomes to compile an all-inclusive dataset with 391 terminals and a reduced dataset with 61 terminals representing main families of Curculionoidea and subfamilies within Curculionidae. Our maximum likelihood and Bayesian phylogenetic analyses, employing both DNA and amino acids datasets under alternative partition schemes, consistently produced congruent phylogenies. Our results show that the Mediterranean raymondionymines form a strongly supported clade, and their easternmost and morphologically distinct genus Ubychia is sister to the rest of them. Most notably, our results consistently recover a sister relationship between the clade of Mediterranean raymondionymine weevils and a clade encompassing all remaining Curculionidae. Consequently, we propose a revision of weevil taxonomy: (i) Our target group is removed from the non-monophyletic subfamily Brachycerinae; (ii) this clade is resurrected to its former subfamily level within Curculionidae, as the subfamily Raymondionyminaestat. rev; (iii) the nine Mediterranean genera Alaocephala, Alaocyba, Coiffaitiella, Derosasius, Ferreria, Raymondiellus, Raymondionymus, Tarattostichus, and Ubychia compose Raymondionyminaestat. rev; (iv) and non-Mediterranean genera Alaocybites, Bordoniola, Gilbertiola, Homosomus, Neoubychia, and Schizomicrus are considered as “incertae sedis” pending further phylogenetic corroboration. We hypothesize that the remaining Brachycerinae and the non-Mediterranean representatives within Raymondionyminae constitute a series of species-poor early-diverging lineages representing currently unrecognized subfamilies of Curculionidae.

We present the findings of a field survey conducted to study deep soil (=endogean) beetles in Chile. We have analysed 50 soil samples, each weighing about 120 kg, collected from 15 central Chilean forested localities along or near the Coastal Range between Valdivia and Santiago. We illustrate all 2,037 adult specimens of Chilean endogean beetles. These specimens include representatives of 11 families: Carabidae, Corylophidae, Curculionidae (subfamilies Cossoninae, Cyclominae, and Raymondionyminae), an unidentified family of Elateroidea, as well as Eupsilobiidae, Leiodidae, Ptiliiidae, Scarabaeidae, Staphylinidae (subfamilies Aleocharinae, Euaesthetinae, Leptotyphlinae, Osoriinae, Pselaphinae, Scydmaeninae, and Staphylininae), Tenebrionidae, and Zopheridae. The most abundant groups are Leptotyphlinae, Leiodidae and Pselaphinae accounting for 45%, 17%, and 17% of all specimens, respectively. The number of adult endogean beetles per soil sample varied between 1 and 150, with an average of 41 specimens (equivalent to an average of 0.34 specimens per kilogram of soil). For each of the 15 sampled localities, specimens were classified to morphospecies, and for each morphospecies we provide the most accurate identification currently possible, considering that most of the species detected are either unnamed or belong to taxonomically challenging groups. In addition, for each morphospecies and locality at least one specimen was selected to be individually vouchered, DNA extracted, and sequenced for the reference barcode DNA fragment in Metazoa. A total of 190 specimens were selected, obtaining barcode sequences for 187 (98.4% success rate) which are provided in an open access DNA barcode library at BOLD Systems (dx.doi.org/10.5883/DS-VGDS28). Obtained sequences were clustered by BOLD Systems in 119 BINs, all corresponding to unique BINs exclusively formed by the newly generated sequences. In addition, when compared to NCBI databases, sequences yielded no matches with a similarity higher than 97%. The comparison with both, BOLD Systems and NCBI databases, supports that none of the deep soil beetle species here collected in Chile has been previously DNA barcoded. We conclude that the deep-soil beetle fauna of Chile is much richer than previously known and mainly unnamed at the species level.

The subfamily Phimophorinae includes ~113 species of cryptic assassin bugs found in the Neotropics and Paleotropics. Presumably due to its small size, cryptic coloration, and occurrence in the remote Eastern Arc Mountains (EAM), Porcelloderes impenetrabilis Rédei, 2012 (Heteroptera: Reduviidae: Phimophorinae), was only recently described as the first representative of this subfamily from mainland Africa. Previous morphology-based phylogenetic research on Phimophorinae (then known as Physoderinae or Epiroderinae) recovered Porcelloderes as sister taxon to the remaining epiroderine-lineage of Phimophorinae. Recent fieldwork in Tanzania has resulted in 241 additional specimens of Porcelloderes from 3 EAM) ranges. We used molecular and morphological analyses for these 241 specimens to (i) determine their species identity, (ii) test the phylo-genetic position of Porcelloderes within Phimophorinae, and (iii) determine if divergence within Porcelloderes coincides with periods of climate-driven forest fragmentation in the EAM. Specimens from the Kimboza and Nguru Mountains likely represent P. impenetrabilis, but we here describe Porcelloderes harles, n. sp., from the Udzungwa Mountains. Our molecular phylogeny shows Porcelloderes nested within the epiroderine-lineage of Phimophorinae and as sister taxon to a Madagascar clade. We estimate that the epiroderine-lineage diverged from other Phimophorinae ~44 MYA. The divergence estimates for the 2 Porcelloderes species is ~15 MYA, consistent with proposed dates for other EAM assassin bugs and a period of forest fragmentation.

The Eastern Arc Mountains (EAMs) in Tanzania and Kenya are renowned for their exceptional species endemism often restricted to isolated mountain blocks. Forest fragmentation during the Pliocene-Pleistocene played a significant role in shaping the EAMs' biodiversity by facilitating allopatric speciation between different mountains. However, only a few studies focused on species diversification within the same mountain block. In this article, we investigated the taxonomy and evolution of the dung beetle genus Grebennikovius, endemic to the Uluguru Mountains. Our goal was to unravel factors promoting allopatric speciation within a confined geographic area like the Ulugurus. We used an integrative taxonomic approach, combining phenotypic data and COI barcodes to delimit species and reconstructed a timetree of the genus. Molecular and morphological evidence consistently recovered 4 distinct Grebennikovius species, of which 3 are new to science: G. armiger n. sp., G. basilewskyi (Balthasar, 1960), G. lupanganus n. sp., and G. pafelo n. sp. Grebennikovius represents a rare case of microal-lopatry, with all species occurring in close proximity within different regions of the limited Uluguru forest (∼230 km 2). We infer that speciation was driven by the interplay between climate fluctuations during the Miocene and mid-Pleistocene, Uluguru topography, and the stability of rainforest ecological conditions. The latter factor probably favored the loss of wings and a specialization for Uluguru microhabitats, reinforcing population isolation and divergence. This study highlights a unique instance of diversification resulting from small-scale speciation events, providing valuable insights into the evolution of Eastern African rainforest arthropods and underscoring the importance of microhabitats conservation.

We present a summary of the geographic distribution data of all 3,341 species of the jewel-beetle genus Agrilus taxonomically valid at the end of 2022. Our work is richly illustrated with maps and diagrams. The genus Agrilus is the most speciose in the Americas (1,292 species) and Asia (1,187 species). The Americas have the best-delimited fauna at the regional level, with 100% of Agrilus species unique to the region, followed by those of Africa (95.8%) and Asia (93.4%). Europe has the lowest percentage of unique Agrilus species on all levels (regional, subregional and country). Among the subregions, the genus Agrilus is the most speciose in South-Eastern Asia (784 species), South America (769), Central America (427) and Eastern Asia (342); on the other hand, Micronesia has a single species, Polynesia has six species and the Caribbean has 18 species. New Zealand has no Agrilus. Brazil (477 species), Mexico (317) and Laos (265) are the three countries with the highest number of Agrilus species. Among 3,341 Agrilus species, 2,924 (87.5%) are unique to one biogeographic realm or transitional zone, and 417 species (12.5%) are shared by two or more biogeographical realms or transitional zones. Three widely distributed Agrilus species found in four biogeographic realms or transitional zones (A. acutus, A. auriventris and A. occipitalis) are pests of cultivated plants Citrus, Abelmoschus, Corchorus, Hibiscus, Malachra, Urena, are likely native to Southeastern Asia and were dispersed by humans. The number of Agrilus species by biogeographic realm varies between 12 (Oceanic) and 1,115 (Neotropical). The genus Agrilus is most speciose in the following realms: Neotropical (1,115 species, 33.4%), Indomalayan (810, 24.2%) and Afrotropical (694, 20.8%). The percentage of unique Agrilus species by biogeographic realm or transitional zone varies between 15.2% (Nearctic–Neotropical transitional zone) to 95.7% (Afrotropical realm). The range of 2,803 Agrilus species (83.9%) lies within a single biogeographic realm: Neotropical (974 species), Afrotropical (665), Indomalayan (662), Palearctic (234) Australasian (174), Nearctic (86) and Oceanic (8). Agrilus species reported from the largest number of countries are A. viridis, A. cuprescens, A. pratensis, A. angustulus, A. biguttatus, A. subauratus, A. cyanescens, A. suvorovi, A. hyperici and A. graminis; they are all native to the Palearctic realm. French Guiana (1.81), Laos (1.12), Vietnam (0.54) and Malaysia (0.53) are the countries with the highest number of Agrilus species per thousand square kilometers. On the sub-country level, the Laotian provinces Houaphan (7.09) and Bolikhamxai (5.52), as well as the Thai province Mae Hong Son (6.55) have the highest number of Agrilus species per thousand square kilometers. Indochina, French Guiana and Mexico are the global hotspots of Agrilus diversity. Our estimates suggest that the true number of Agrilus species in subtropical and tropical zones is at least twice higher than is currently known.

We use DNA sequence data to generate the first phylogenetic hypothesis for the weevil tribe Lymantini. These are leaf litter inhabiting beetles generally regarded as restricted to the New World and taxonomically arranged in two subtribes, 11 genera and some 150 named species. An additional genus of questionable affinities to the tribe, Devernodes Grebennikov, 2018, has five described species in Southeastern Asia. All these beetles are flightless and some have eyes reduced in size or absent, traits normally associated with limited dispersal capacity. We performed a phylogenetic analysis of 153 terminals (50 of them belong to Lymantini representing Devernodes and all but three named genera) based on 4,174 bp alignment of one mitochondrial (cox1) and two nuclear fragments (ITS2 and 28S). We find that both Lymantini subtribes Lymantina and Caecossonina are monophyletic, the latter sister to the amphi-Atlantic tribe Anchonini. The Asian genus Devernodes is deeply nested among American Lymantina. The clade of Anchonini plus Lymantini is consistently recovered outside of the CCCMS clade of “higher” weevils (Curculioninae, Conoderinae, Cossoninae, Molytinae and Scolytinae). We hypothesize that the polished head capsule of adult beetles is an apomorphy of Anchonini and Lymantini, the 8-segmented antennal funicle is an apomorphy of Anchonini plus Caecossonina. We attribute the origin of the currently observed amphi-Pacific distribution of Lymantina to normal ecological dispersal facilitated by the warmer periods of the Cenozoic such as the Eocene, and by presently submerged Arctic land bridges. Using parsimony we hypothesize a North American origin for the Anchonini plus Lymantini crown group, as well as that of Lymantina. We argue that Bronchotibia adunatus Poinar & Legalov, 2021, a Dominican amber adult weevil fossil, is not a member of Lymantini and re-classify it as Curculionidae incertae sedis. We present an image gallery of 28 Lymantini specimens to document the morphological diversity of the tribe. We hypothesize the existence of unnamed American genera of Lymantina and make public the DNA-barcode dataset of 89 Lymantini specimens.

Wir klären die taxonomische Zugehörigkeit und die phylogenetische Stellung zweier konspezifischer weiblicher Rüsselkäfer aus der Streuschicht des Huisun-Waldes auf Taiwan. Diese Rüsselkäfer sind denjenigen der Gattungen Protacalles Voss, 1957, Protacallinus Morimoto, 1962 und Protacallodes Morimoto, 2011 am ähnlichsten, welche alle endemisch für Japan sind. Wir führten eine phylogenetische Analyse mit 37 Taxa und 83 parsimonieinformativen morphologischen Merkmalen durch. Der sparsamste Baum ergab eine Klade dieser drei Gattungen, mit der Art aus Taiwan als Schwesterart zu Protacallodes. Der Baum wird statistisch nur schwach unterstützt und die Kladen kollabierten nach dem Bootstrapping-Verfahren. Wir beschreiben die taiwanesischen Rüsselkäfer als Protacallodes bhudevi sp. n. Dies ist die dritte Art der Gattung und der Erstnachweis der Gattung für Taiwan. Die neue Art und ihre möglichen verwandten Arten werden illustriert und in einem angepassten Bestimmungsschlüssel vorgestellt.

Paleotropical clades with largely disjunct distributions are ideal models for biogeographic reconstructions. The dung beetle genera Grebennikovius Mlambo, Scholtz & Deschodt, Epactoides Olsouffief and Ochicanthon Vaz‐de‐Mello are distributed in Tanzania, Madagascar and Réunion, and the Oriental region, respectively. We combine morphology and molecular dataset to reconstruct the phylogenetic relationships between these taxa. Our analyses corroborate previous hypotheses of monophyly of the group, which is here described as new tribe Epactoidini trib. nov. Grebennikovius is recovered as sister to Epactoides, while Ochicanthon emerges as sister to them both. The disjunct distribution of our focal clade is unusual within the subfamily Scarabaeinae. Bayesian divergence time estimates and ancestral range reconstructions indicate an African origin of the crown group of the tribe Epactoidini trib. nov. in the early mid Eocene, ca. 46 Ma. The divergence between Epactoides and its sister is dated to 32.3 Ma, while the crown age for the genus Ochicanthon is dated to 27 Ma. We investigate the factors that may have shaped the current distribution of the tribe Epactoidini trib. nov. The formation of the Gomphotherium landbridge, along with favourable environmental conditions would have allowed dry‐intolerant organisms, such as Ochicanthon, to disperse out of Africa. Remarkable climatic stability of the Eastern Arc Mountains was critical for the retention of the monotypic genus Grebennikovius. We suggest two subsequent overwater dispersal events: the migration of the most recent common ancestor (MRCA) of Epactoides from Africa to Madagascar (32.3–29.5 Ma); the lately dispersal of the MRCA of the today's extinct Epactoides giganteus Rossini, Vaz‐de‐Mello & Montreuil to Réunion island from Madagascar (3.4 Ma). We suggest that the high potential of dispersal of Epactoidini trib. nov. dung beetles and the strict association to forest habitat might have triggered two major radiations, one in Madagascar and one in the Oriental Region. This study corroborates the monophyly of the dung beetle clade Grebennikovius + Epactoides + Ochicanthon, which is described as a new tribe Epactoidini trib. nov. Biogeographic reconstructions indicate an African origin for Epactoidini (ca 46 Ma). Ochicanthon would have dispersed to Asia through the Gomphotherium landbridge (ca 46–27 Ma). Eastern Arc Mountains played as retention area for Grebennikovius, while Epactoides would have reached Madagascar (32.3–29.5 Ma) and Réunion island (3.4 Ma) through two overwater dispersal events.

Paleotropical clades with largely disjunct distributions are ideal models for biogeographic reconstructions. The dung beetle genera Grebennikovius Mlambo, Scholtz & Deschodt, Epactoides Olsouffief and Ochicanthon Vaz-de-Mello are distributed in Tanzania, Mada-gascar and Réunion, and the Oriental region, respectively. We combine morphology and molecular dataset to reconstruct the phylogenetic relationships between these taxa. Our analyses corroborate previous hypotheses of monophyly of the group, which is here described as new tribe Epactoidini trib. nov. Grebennikovius is recovered as sister to Epactoides, while Ochicanthon emerges as sister to them both. The disjunct distribution of our focal clade is unusual within the subfamily Scarabaeinae. Bayesian divergence time estimates and ancestral range reconstructions indicate an African origin of the crown group of the tribe Epactoidini trib. nov. in the early mid Eocene, ca. 46 Ma. The divergence between Epactoides and its sister is dated to 32.3 Ma, while the crown age for the genus Ochicanthon is dated to 27 Ma. We investigate the factors that may have shaped the current distribution of the tribe Epactoidini trib. nov. The formation of the Gomphotherium landbridge, along with favourable environmental conditions would have allowed dry-intolerant organisms, such as Ochicanthon, to disperse out of Africa. Remarkable climatic stability of the Eastern Arc Mountains was critical for the retention of the monotypic genus Grebennikovius. We suggest two subsequent overwater dispersal events: the migration of the most recent common ancestor (MRCA) of Epactoides from Africa to Madagascar (32.3-29.5 Ma); the lately dispersal of the MRCA of the today's extinct Epactoides giganteus Rossini, Vaz-de-Mello & Montreuil to Réunion island from Madagascar (3.4 Ma). We suggest that the high potential of dispersal of Epactoidini trib. nov. dung beetles and the strict association to forest habitat might have triggered two major radiations, one in Madagascar and one in the Oriental Region.

This paper targets diversity and phylogeny of the Old World weevil tribe Blosyrini and, specifically, its Asian members. Phylogenetic analysis of one mitochondrial and two nuclear DNA fragments from 78 terminals reveals that Blosyrini weevils, although monophyletic in Asia, in Madagascar, and in continental Africa, do not share a unique common ancestor. Instead, they form a strongly supported clade together with representatives of two other tribes of broad-nosed weevils: Cneorhinini and Dermatodini. The Asian members of the tribe form a moderately supported clade, of which the monophyletic genus Trachyphloeoides is a sister to the strongly supported rest, Blosyrini Clade X (BCX). Owing to the convoluted and non-phylogenetic taxonomy, BCX cannot be at present reliably referred to by any existing genus-group name. All 112 DNA barcodes of BCX (including one larva) from China, Kazakhstan, Kyrgyzstan, and Nepal form 34 Barcode Index Numbers (BINs). Each of seven comprehensively sampled mountainous localities in Sichuan (Gongga Shan, Emei Shan, Songpan) and Yunnan (Cang Shan, Gaoligong Shan, Haba Shan, Jizu Shan) supports between one and six BINs of BCX. With two exceptions, all BINs of BCX in Sichuan (8) and Yunnan (10) display strong biological preferences for either mid-altitude primary deciduous forests or the high elevation alpine zone. Seven strongly supported clades are recognized within BCX, some of them morphologically diagnosable. Temporal analysis corroborates the results of BIN clustering and interrelationships within BCX. The most recent common ancestor of BCX lived in the mid-Miocene (14.15 Ma, 95% confidence interval 17.5–11.2 Ma), with much of the subsequent diversification preceding or coinciding with the Pliocene-Pleistocene climatic fluctuations.