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Recent phylogenetic studies in Himalayan lazy toads of the genus Scutiger Theobald, 1868 revealed the presence of genetically deeply divergent lineages. The taxonomy of S. nepalensis sensu lato was re-assessed based on museum material considering molecular and morphological data. The results strongly support the recognition of a new species, S. kanjiroba sp. nov. distributed along the Nepalese Kanjiroba massif. It is further shown that S. sikimmensis has an apparently much more restricted distribution range than previously thought. The frequent misidentification of Scutiger across large areas of the Himalaya-Tibet area highlights the significance of careful taxonomic evaluation of collection material and the need for the direct morphological comparison of closely related species when describing new species.

The trechine beetle fauna (Coleoptera, Carabidae) of the Ethiopian Highlands is known to be highly diverse in species, and many species groups were recognized to be characterized by unusual character states of external and genital morphology. Earlier authors described several genera and subgenera of Ethiopian Trechina endemic to certain high mountains of the country. However, the relationships of these species groups and their evolutionary history are unknown so far. Here, we present the first molecular phylogenetic analysis of Ethiopian Trechina, detect several synonymic names under Trechus sensu lato, and introduce two new species groups to the country’s fauna: the monotypic genus Baehria Schmidt & Faille, gen. nov., with the type species B. separata sp. nov. from Mt. Choke in northern Ethiopia, and the Trechus subgenus Abunetrechus Schmidt & Faille, subgen. nov., with the type species T. bipartitus Raffray, 1885; this subgenus includes three species of northern Ethiopia. We show that the composition of the Ethiopian fauna is based on multiple events of immigration, which started simultaneously with or some million years after the Oligocene-Early Miocene orogenic events north and south of the Rift Valley. Our results support the habitat island hypothesis for the evolution of the Ethiopian highland fauna. We found no evidence for an alternative hypothesis assuming a close connection of the Trechina immigration to Ethiopia and Pleistocene cooling. We, thus, conclude that the geomorphological development rather than the climatic changes are the main drivers of the diversification of the high-altitude Trechina fauna in Ethiopia.

Of the 12 specimens of Calathus-like sphodrine beetles presently known from Baltic and Rovno amber deposits, 11 specimens were investigated using light microscopy, micro-CT scanning, and 3D imaging techniques. For the first time, many significant diagnostic characters of the external morphology and male and female genitalia of Eocene Sphodrini were studied in detail. Based on these data, three fossil species are diagnosed and placed in a natural group characterized by a derived pattern in elytral chaetotaxy and microsculpture and therefore the genus Quasicalathus Schmidt & Will, gen. nov. is described to comprise these species. Due to the presence of a styloid right paramere, Quasicalathus gen. nov. is considered a member of the sphodrine “P clade” of Ruiz et al. (2009). However, given the absence of synapomorphies of any species group of the P clade, the systematic position of Quasicalathus gen. nov. within this clade remains unresolved. The Baltic amber species Calathus elpis Ortuño & Arillo, 2009 is redescribed based on additional, fossil, non-holotype material and transferred to Quasicalathus gen. nov. Identification of the additional C. elpis fossil material remains slightly uncertain due to the non-availability of the holotype for direct comparison coupled with doubts regarding the accuracy of certain character states presented in its original description. Two species are newly described: Quasicalathus agonicollis Schmidt & Will, sp. nov., from Baltic amber, and Q. conservans Schmidt & Will, sp. nov., from Rovno amber.

Balticeler kerneggeri gen. nov., sp. nov., is described based on six fossil specimens preserved in Eocene Baltic amber and imaged using light microscopy and X-ray micro-computed tomography. Based on certain characters observed in the fossil species it is considered a “middle grade” Carabidae, outside of the large family Harpalinae (as it possesses a scrobal seta, the lack of which is a synapomorphy of that subfamily), but possessing four synapomorphies that indicate Balticeler belongs to a large clade of carabids including Harpalinae (anisochaetous Grade B antennal cleaner, conjunct mesocoxae, closed procoxal cavities, and a well-developed external lobe of the metepimeron). This remarkable beetle has several striking features, including lack of externally-visible sexually dimorphic characters, lack of lateral borders on the pronotum, and very long and thin mandibles and maxillae. In combination, these states are unique within Carabidae. We consider the presence of a dorsally completely open aedeagal median lobe as a synapomorphy of the fossil species with the subfamily Trechinae, a pubescent and relatively long second antennomere and a 4+2+2 pattern of umbilicate setae as synapomorphies of the supertribe Trechitae, and a quadrisetose clypeus as a synapomorphy with the Trechitae clade Bembidarenini + Trechini sensu Maddison et al. (2019). As it lacks a synapomorphy of Bembidarenini + Trechini, we propose that it is a member of the stem group of that clade.

The phylogeny of the carabid beetle supertribe Nebriitae is inferred from analyses of DNA sequence data from eight gene fragments including one nuclear ribosomal gene (28S), four nuclear-protein coding genes (CAD, topoisomerase 1, PEPCK, and wingless), and three mitochondrial gene fragments (16S + tRNA-Leu + ND1, COI (“barcode” region) and COI (“Pat/Jer” region)). Our taxon sample included 264 exemplars representing 241 species and subspecies (25% of the known nebriite fauna), 39 of 41 currently accepted genera and subgenera (all except Notiokasis and Archileistobrius), and eight outgroup taxa. Separate maximum likelihood (ML) analyses of individual genes, combined ML analyses of nuclear, nuclear protein-coding, and mitochondrial genes, and combined ML and Bayesian analyses of the eight-gene-fragment matrix resulted in a well-resolved phylogeny of the supertribe, with most nodes in the tree strongly supported. Within Nebriitae, 167 internal nodes of the tree (out of the maximum possible 255) are supported by maximum-likelihood bootstrap values of 90% or more. The tribes Notiophilini, Opisthiini, Pelophilini, and Nebriini are well supported as monophyletic but relationships among these are not well resolved. Nippononebria is a distinct genus more closely related to Leistus than Nebria. Archastes, Oreonebria, Spelaeonebria, and Eurynebria, previously treated as distinct genera by some authors, are all nested within a monophyletic genus Nebria. Within Nebria, four major clades are recognized: (1) the Oreonebria Series, including eight subgenera arrayed in two subgeneric complexes (the Eonebria and Oreonebria Complexes); (2) the Nebriola Series, including only subgenus Nebriola; (3) the Nebria Series, including ten subgenera arrayed in two subgeneric complexes, the Boreonebria and Nebria Complexes, with the latter further subdivided into three subgeneric subcomplexes (the Nebria, Epinebriola, and Eunebria Subcomplexes)); and (4) the Catonebria Series, including seven subgenera arrayed in two subgeneric complexes (the Reductonebria and Catonebria Complexes). A strong concordance of biogeography with the inferred phylogeny is noted and some evident vicariance patterns are highlighted. A revised classification, mainly within the Nebriini, is proposed to reflect the inferred phylogeny. Three genus-group taxa (Nippononebria, Vancouveria and Archastes) are given revised status and seven are recognized as new synonymies (Nebriorites Jeannel, 1941 and Marggia Huber, 2014 = Oreonebria Daniel, 1903; Pseudonebriola Ledoux & Roux, 1989 = Boreonebria Jeannel, 1937; Patrobonebria Bänninger, 1923, Paranebria Jeannel, 1937 and Barbonebriola Huber & Schmidt, 2017 = Epinebriola Daniel & Daniel, 1904; and Asionebria Shilenkov, 1982 = Psilonebria Andrewes, 1923). Six new subgenera are proposed and described for newly recognized clades: Parepinebriola Kavanaugh subgen. nov. (type species: Nebria delicata Huber & Schmidt, 2017), Insulanebria Kavanaugh subgen. nov. (type species: Nebria carbonaria Eschscholtz, 1829), Erwinebria Kavanaugh subgen. nov. (type species Nebria sahlbergii Fischer von Waldheim, 1828), Nivalonebria Kavanaugh subgen. nov. (type species: Nebria paradisi Darlington, 1931), Neaptenonebria Kavanaugh subgen. nov. (type species: Nebria ovipennis LeConte, 1878), and Palaptenonebria Kavanaugh subgen. nov. (type species: Nebria mellyi Gebler, 1847). Future efforts to better understand relationships within the supertribe should aim to expand the taxon sampling of DNA sequence data, particularly within subgenera Leistus and Evanoleistus of genus Leistus and the Nebria Complex of genus Nebria.

The ground beetle genus Bembidion is a highly diverse group of small predators with more than 1.200 described extant species. In contrast, only two representatives of Bembidion are known from the amber fossil record and their position within this mega-diverse genus is dubious. Here, we address the taxonomic position of these two extinct Bembidion species (B. succini Giebel, 1856 and B. christelae Ortuño & Arillo, 2010). Based on the insufficient description and the missing type specimen, B. succini, nomen dubium, cannot be assigned to the genus Bembidion and/or to the tribe Bembidiini with certainty. The subgenus Archaeophilochthus Ortuño & Arillo, 2010 was erected for the second extinct species, B. christelae, based on external characters. However, this species seems indistinguishable to members of the earlier described subgenus Philochthemphanes Netolitzky, 1943 which comprises about extant 10 species distributed in East and Southeast Asia. Furthermore, we describe two new species, B. bukejsi sp. n. and B. alekseevi sp. n., from the Eocene Baltic amber using X-ray microscopy. Based on external and genital morphology including endophallic structures, we erected the monotypic subgenus Eodontium subgen. n. for B. bukejsi sp. n., which is probably related to the subgenera Andrewesa Netolitzky, 1931, the Hydrium complex, or the Odontium series sensu Maddison (2012). On the other hand, B. alekseevi sp. n. can be assigned to the subgenus Eupetedromus Netolitzky, 1911. The occurrence of representatives of at least two species groups adapted to a temperate climate suggests the presence of at least locally temperate climates in Baltic amber forests.